Deep-Sea Research II ∎ (∎∎∎∎) ∎∎∎–∎∎∎

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Deep-Sea Research II
journal homepage: www.elsevier.com/locate/dsr2

Introduction

Towards a new and integrated approach to submarine canyon research

art ic l e i nf o

Keywords:
Submarine canyons
International network
Multidisciplinary research
Geology
Biology
Oceanography

1. Background increasing availability of new sampling and surveying technologies

(deep-towed acoustic instruments, drop-down video systems, and
Submarine canyons, steep-walled valleys that cut across virtu eventually robotic vehicles), submarine canyon research increased
ally every continental margin around the world (Harris and dramatically (Fig. 1). Particularly the advent of Remotely Operated
Whiteway, 2011), are considered major sediment transport path- Vehicles (ROVs) in many research institutes in the last
10 years
ways between continental shelves and the deep sea (e.g., Shepard, opened up a new perspective on submarine canyons, allowing a
1963; Puig et al., 2014). Owing to their steep topography and high wider community of researchers to access parts of the deep ocean
terrain heterogeneity, in addition to their unique current patterns that had been hidden until then (Tyler et al., 2009).
and episodic down-canyon flushing events, which result in locally As a result of this increased research effort, our understanding
increased nutrient concentrations and food availability, submarine of submarine canyons is gradually growing. A number of indivi-
canyons are often considered as biodiversity hotspots (e.g., Tyler dual canyon systems have received considerable attention (e.g. the
et al., 2009; De Leo et al., 2010). On the other hand, consider- Portuguese Canyons offshore Lisbon & Nazaré (Masson and Tyler,
able differences have been observed between individual canyon 2011 and references therein); Monterey Canyon (e.g., Hall and
systems, and between different faunal groups in terms of their Carter, 2011; Paull et al., 2011; Robison et al., 2010) or the Cap
response to the typical canyon environment (e.g., Cunha et al., de Creus Canyon and the other canyons in the Gulf of Lions
2011; Ingels et al., 2011; Schlacher et al., 2007). Unfortunately, in (e.g., Canals et al., 2006; Lastras et al., 2007; Orejas et al., 2009;
addition to transporting sediment, submarine canyons also tend to Palanques et al., 2008)) but most canyons around the world have
funnel our human litter and pollutants into the deep sea, extend- not yet been studied, or only to a very limited extend. Further-
ing the anthropogenic impact on the oceans far beyond our shores more, many of the studies carried out so far are focussing on
(e.g., de Jesus Mendes et al., 2011; Mordecai et al., 2011; Schlining one aspect (geology, geomorphology, sediment dynamics, hydro-
et al., 2013). graphy, current patterns, mega-, macro-, meiofauna distribution,
Submarine canyons have been the subject of research for a long biogeochemistry…) of a single canyon or canyon system. The time
time. Shepard (1972) refers to a study from as early as the late seems right to start putting all those pieces of the jigsaw together,
nineteenth century, carried out by Milne (1897), which looked and to start looking at canyons in a more holistic way. To this end,
at the instability of canyon floor sediments as a possible cause for the first International Symposium on Submarine Canyons was
the repeated breaking of submarine cables that had been laid organised in Brest, France in July 2012. Canyon research from all
across a canyon. However, as a result of the steep terrain, locally over the world was presented, followed by cross-disciplinary
enhanced currents and occasional down-canyon flushing events, discussions and networking. A good proportion of those studies
the initial submarine canyon investigations were extremely chal- are presented here, in this Special Issue. In addition, the meeting
lenging, and the number of studies was limited. Acoustic methods resulted in the formation of INCISE: the International Network for
had to deal with excessive scatter and noise, in-situ instruments submarine Canyon Investigation and Scientific Exchange (www.
were regularly washed away and the coarse canyon thalwegs and incisenet.org). Further meetings and sessions at international
rocky walls proved difficult to sample (e.g., Paull et al., 2003; conferences are planned, and an active forum has been set up,
Shepard, 1972). Direct observations were limited to shallow with the aim to stimulate cross-disciplinary discussions and
waters, within reach of divers or early submarines. With the research activities.

0967-0645/$ - see front matter & 2013 Published by Elsevier Ltd.

http://dx.doi.org/10.1016/j.dsr2.2013.09.012
2 Introduction / Deep-Sea Research II ∎ (∎∎∎∎) ∎∎∎–∎∎∎

2. Integrated submarine canyon research The upper waters (o750 m) are dominated by species typical of the
mid- to higher latitudes in the N. Atlantic, which spend part of their
This Special Issue presents submarine canyon research from all daily cycle in the surface waters brought in by the overall NE–SW
over the world (Fig. 2). Particularly the large number of studies on the current. Inter-annual variability in species abundance is limited,
Sable Gully, off Nova Scotia, illustrates the type of integrated picture however, despite substantial changes in oceanographic conditions
INCISE hopes to achieve for many individual canyons, and for during the 3-year study by MacIsaac et al. (2013). In contrast,
submarine canyons as a whole. The Sable Gully was the first Canadian Kenchington et al. (2013) looked at the epibenthic macrofauna living
Marine Protected Area (MPA) to be designated in the Atlantic. The area in the tidally dominated deeper parts of the canyon, beyond 1000 m
is known to be an important cetacean habitat; especially northern depth. They found that those fauna mainly consist of filter-feeders,
bottlenose whales seem to have a great affinity for this canyon and which probably reflects the influence of the tidal currents in the
similar canyons in the region (Moors-Murphy, 2013). Oceanographic/ canyon. Furthermore, apart from depth, benthic communities are also
hydrographic observations by Greenan et al. (2013) illustrate the structured by food availability (total organic carbon and labile carbon)
unique tidal environment of the canyon, which is dominated by at a spatial scale of 10 s of km, while on shorter distances (10 s of m)
unusual non-linear constituents that create overtides and compound substratum seems to be an important factor determining species
tides. In addition, there is strong evidence for enhanced mixing and associations.
up-canyon flow within the canyon. The surface waters, however, do Of course, as indicated before: patterns emerging from a single
not seem to be very much affected by the presence of the canyon: they canyon system may not necessarily be universal. Comparing
are mainly influenced by the regional NE–SW current pattern. To canyons located on the active transform margin off Haida Gwaii,
further enhance our understanding of the 3-dimensional circulation British Columbia, with the Sur Canyon system on the other well-
throughout the canyon, Shan et al. (2013) applied a multi-nested known transform margin south of Monterey Bay, California, Harris
ocean circulation model, investigating the influence of shelf-scale et al. (2013) demonstrate that several factors are at play in the
circulation, tide-topography interaction and wind forcing on the creation of canyon morphologies. Considerable variability has also
circulation within and above the canyon. The authors found that been found between canyons as close as a few 10s–100s of
wind, especially during storm events, is a significant factor affecting kilometres apart. For example, Lo Iacono et al. (2013) present
the circulation above the canyon, while especially the tide-topography two submarine canyons located offshore N Sicily. Although both
interaction is a dominant factor within the Gully. This water mass are placed within the same tectonically active geological setting,
structure and current pattern may well affect the pelagic fauna. For the canyons have a very different morphology, and are governed
example, the crustacean micronekton and macrozooplankton are by different sedimentary processes. Bottom-up, retrogressive slope
mainly structured by depth and diel cycle (MacIsaac et al., 2013). failures have driven the formation of the Palermo Canyon, while
top-down, erosive turbidity currents, linked to fluvial sedimentary
input, have shaped the meandering Castellamare Canyon. Similar,
although less extreme, differences were found by Obelcz et al.
(2013) in the fine-scale morphology of four submarine canyons,
spaced over 200 km along the US Mid-Atlantic passive margin,
pointing to slightly different levels of canyon activity and sediment
transport processes.
However, in terms of recent, short-term sediment transport in
shelf-incising canyons, most observations seem to confirm the
overall two-step mechanism already hinted by Shepard (1963),
and recently reviewed by Puig et al. (2014). For example, tidally
mobilised shelf sediments are intercepted by the Cook Strait
Canyon heads offshore New Zealand, and accumulate in depocen-
tres in the upper/central part of the canyon system. They are then
Fig. 1. Number of publication records in the Web of Knowledge database related to
remobilised in more catastrophic events and transported towards
the search topic ‘submarine canyon’ as of 5 August 2013. the lower canyon and deep ocean as a result of tectonic (earth-
Source: Thomson Reuters. quake) processes with a return period of ca. 100 years (Mountjoy

Fig. 2. Locations of submarine canyon studies published in this Special Issue. Labels refer to the order of the papers in this journal.
 Introduction / Deep-Sea Research II ∎ (∎∎∎∎) ∎∎∎–∎∎∎ 3

et al., 2013). Similarly, the middle Cap-Ferret Canyon (Bay of in the discovery of new species (e.g. Kenchington et al., 2013;
Biscay, N Atlantic) acts as a depocentre for material that has been MacIsaac et al., 2013; Schejter et al., 2013). It will be clear that
intercepted from outer shelf and upper slope transport by the much more information is needed, including a deeper under-
canyon head and has bypassed the upper canyon. The lower standing of canyon biology and ecology, before policy-makers will
canyon only receives sediment input in the form of gravity flows be able to design the optimal submarine canyon management
on timescales of decades or more (Duros et al., 2013; Schmidt plans and conservation measures. Integrating the range of biolo-
et al., 2013). gical studies presented in this Special Issue (in addition to the
On even shorter timescales, observations in countless submar- wider submarine canyon literature), some patterns begin to
ine canyons (including the work by Greenan et al. (2013) and emerge, although a lot of uncertainty still exists about the details.
Shan et al. (2013) in the Gully) have shown that internal tides and It is obvious that, as reported by Kenchington et al. (2013) and
waves are some of the most important factors in the daily (re)- MacIsaac et al. (2013) for the Sable Gully, depth is the main
suspension and flux of canyon sediments. Hall et al. (2013) now structuring factor in the distribution of most canyon fauna/groups
demonstrate that the nature of this internal tidal driving force can (Duffy et al., 2013; De Leo et al., 2013; Frutos and Sorbe, 2013).
change from a standing to a progressive wave as the result of a However, further driving factors remain difficult to ascertain. Duffy
change in watercolumn stratification. Using a numerical model of et al. (2013) could not find a statistically significant difference
Monterey canyon, they show that an observed change in depth of between the epibenthic megafauna communities of 5 canyons
the main pycnocline from 200 m (below the canyon rim) to 50 m with different sediment transport regimes offshore California.
(above the canyon rim) decreases the supercritical reflection of the However, the authors admit that the nature of their dataset
up-canyon travelling internal tidal wave, allowing it to become (opportunistic ROV-based video rather than imagery transects
dominant over the reflected down-canyon wave. This affects the conducted according to a rigorous sampling design) may play an
energy flux through the canyon, and the spatial pattern of current important role in this result. This was not an issue in the work of
amplification and sediment resuspension at the seabed. However, De Leo et al. (2013), who sampled infaunal macrobenthos in
in addition to natural processes, there is increasing evidence that 6 Hawaiian canyons, using a depth-stratified scheme. They found
daily sediment resuspension and transport may also be caused that habitat heterogeneity at medium and large spatial scales, in
by anthropogenic activities such as deep-water bottom traw- addition to Particulate Organic Carbon (POC) and distance from
ling. Martin et al. (2013) present a stunning dataset illustrating shore (both proxies for food input) seem to be good predictors of
the daily occurrence of increased water column turbidity and macrobenthic biodiversity. Food availability even seems to be the
the associated sediment gravity flows in La Fonera Canyon (NW main structuring factor in Kaikoura Canyon, offshore New Zealand
Mediterranean). The phenomenon occurs exclusively during the (Leduc et al., 2013). This canyon is characterised by unusually high
working days/hours of the bottom trawling fleet that fishes on food availability for benthic fauna, probably due to a combination
the upper canyon flanks. The process is gradually reshaping the of increased primary production and downwelling/topographically
canyon morphology, but also influences faunal communities in the steered funnelling of surface-derived organic matter. The authors
deeper reaches, by creating increased disturbance and sediment looked at free-living nemathodes in the canyon, and found an
turbidity compared to the natural state. exceptionally high biomass, reduced diversity and specific nema-
Sediment resuspension is not the only type of anthropogenic tode community structure. On a more local scale, disturbance has
impact reported in the submarine canyon studies of this Special also been reported as an influencing factor in submarine canyons.
Issue. Lost fishing gear is very common, as is litter (e.g., Fabri et al., Although less than a kilometre apart, benthic foraminifera in
2013; Davies et al., 2013). Fabri et al. (2013) carried out a compre- samples from the Capbreton Canyon thalweg show a distinctively
hensive study of 17 canyons along the French Mediterranean margin different assemblage than those on the lower canyon flank. They
in order to evaluate the occurrence and status of Vulnerable Marine have a lower diversity and are dominated by pioneer species,
Ecosystems (VMEs) in the area. They found that sea pen grounds illustrating that the community stays in an early stage of ecosys-
(Pennatulacea) and Alcyonacea grounds were fairly rare, and pre- tem colonisation as a result of the regular deposition of turbidite
sented lower faunal densities than expected. This could be related to sequences (Bolliet et al., 2013). Taking an entirely different
trawling pressure. Cold-water corals such as Lophelia pertusa and perspective, Guerreiro et al. (2013) present one of the first studies
Madrepora oculata were not very common, and in addition to to characterise surface water coccolithophore assemblages in the
damage by lost fishing gear, were also affected by the discharge of context of an active submarine canyon and its associated physical
bauxite mud in Cassidaigne Canyon. A similar benthic megafauna oceanography. They find that the waters around Nazaré Canyon,
biotope study was carried out by Davies et al. (2013) in the upper offshore Portugal, are dominated in winter by two assemblages,
part of a submarine canyon system along the Celtic Margin, NE distinguishing slightly more nutrient-rich coastal-neritic waters
Atlantic. They found 12 biotopes, of which 4 can be classified as (influenced by surface water runoff) from the mixed oceanic
VMEs. Especially Kophobelemnon (sea pen) grounds were specific for waters advected onshore by the canyon. In addition, they report
the canyons, but biotopes harbouring L. pertusa were present as well. that the canyon head is often a location of high productivity
In addition, Stewart et al. (2013) report the discovery of large between March and October, creating a coccolithophore diversity
numbers of small mounds in the same area (up to 4 m high and hotspot, mixing both the oligotrophic-oceanic and opportunistic
150 m across, located at water depths of 250–400 m) covered with coastal taxa. Such an increased productivity will have a strong
cold-water coral rubble. It is not clear if the coral died as a result of influence on the benthic environment in the upper-middle Nazaré
natural causes (change in environmental conditions over the last Canyon, where an extremely rich depocentre has been reported at
1000s of years) or of deep-water trawling, which is very intensive ca. 3400 m water depth by several authors (e.g. Amaro et al., 2010;
in the area. Given the struggle of scleractinian corals to survive Cunha et al., 2011; Masson et al., 2010).
anthropogenic impacts in the European canyons, the discovery of
L. pertusa in canyons on the US margin of the N Atlantic is a positive
finding (Brooke and Ross, 2013). 3. Outlook
Although studies like those by Davies et al. (2013) and Fabri
et al. (2013) are invaluable for marine conservation, they only As demonstrated by the case studies of the Sable Gully
cover a small part of the world's oceans and submarine canyons. (Greenan et al., 2013; Kenchington et al., 2013; MacIsaac et al.,
Nearly every biological survey of a (new) submarine canyon results 2013; Moors-Murphy, 2013; Shan et al., 2013), in order to obtain
4 Introduction / Deep-Sea Research II ∎ (∎∎∎∎) ∎∎∎–∎∎∎

a better understanding of submarine canyon systems as a whole, De Leo, F.C., Vetter, E.W., Smith, C.R., Rowden, A.A., McGranaghan, M., this issue.
more integrated research efforts are needed, combining insights Spatial scale-dependent habitat heterogeneity influences submarine canyon
macrofaunal abundance and diversity off the Main and Northwest Hawaiian
on canyon geology, sedimentology and oceanography with obser- Islands. Deep-Sea Research II. [doi: 10.1016/j.dsr2.xxxx.xx.xxx].
vations of biology and ecology. Also human activities and impacts Duffy, G.A., Lundsten, L., Kuhnz, L.A., Paull, C.K., this issue. A comparison of
should be included in the picture (Martin et al., 2013). Deriving megafaunal communities in five submarine canyons off Southern California,
USA. Deep-Sea Research II. this issue [http://dx.doi.org/10.1016/j.dsr2.2013.06.
universal patterns of canyon processes and characteristics
002].
will require such integrated research efforts to be repeated in Duros, P., Jorissen, F.J., Cesbron, F., Zaragosi, S., Schmidt, S., Metzger, E., Fontanier, C.,
several canyons, for several tectonic and environmental settings. this issue. Study of benthic foraminiferal thanatocoenoses from the Cap-Ferret
To achieve comparable results, the submarine canyon research Canyon area (NE Atlantic): A complex interplay between hydro-sedimentary
and biological processes. Deep-Sea Research II. [doi: 10.1016/j.dsr2.xxxx.xx.
community will have to work towards comparable and compatible xxx].
methodologies, including the set-up of (or continuation of) long- Fabri, M.-C., Pedel, L., Beuck, L., Galgani, F., Hebbeln, D., Freiwald, A., this issue.
term monitoring programmes to assess the temporal aspects of Megafauna of vulnerable marine ecosystems in French Mediterranean sub-
marine canyons: spatial distribution and anthropogenic impacts. Deep-Sea
canyon processes (Juniper et al., 2013; Martins et al., 2010). Research II. [doi: 10.1016/j.dsr2.xxxx.xx.xxx].
Understanding the scale (both spatial and temporal) at which Frutos, I., Sorbe, J.C., this issue. Bathyal suprabenthic communities from the
canyon processes shape the canyon environment is key to under- southern margin of the Capbreton Canyon (“Kostarrenkala” fishing ground),
SE Bay of Biscay. Deep-Sea Research II. this issue [http://dx.doi.org/10.1016/j.
standing the biological and ecological patterns. Only when
dsr2.2013.09.010].
we understand the frequency, extent and biological response Greenan, B.J.W., Petrie, B.D., Cardoso, D.A., this issue. Mean circulation and high-
to natural disturbance events, will we be able to assess the real frequency flow amplification in the Sable Gully. Deep-Sea Research II. [doi:
impact of anthropogenic disturbance in this unique environment. 10.1016/j.dsr2.xxxx.xx.xxx].
Guerreiro, C., Sá, C., de Stigter, H., Oliveira, A., Cachão, M., Cros, L., Borges, C.,
Obtaining an insight in the connectivity of submarine canyons will Quaresma, L., Santos, A.I., Fortuño, J.-M., Rodrigues, A., this issue. Late winter
be necessary to devise viable networks of marine protected areas. coccolithophore assemblage from the Nazaré Canyon, central Portuguese
The International Symposium on Submarine Canyons in Brest has margin. Deep-Sea Research II. [doi: 10.1016/j.dsr2.xxxx.xx.xxx].
Hall, R.A., Alford, M.H., Carter, G.S., Gregg, M.C., Lien, R.-C., Wain, D.J., Zhao, Z., this
set in motion a dynamic and ambitious community of researchers
issue. Transition from partly standing to progressive internal tides in Monterey
working towards these goals, aiming to increase the understand- Submarine Canyon. Deep-Sea Research II. [http://dx.doi.org/10.1016/j.dsr2.
ing of submarine canyons in a holistic way. We hope this Special xxxx.xx.xxx].
Issue will be a first step, stimulating further cross-disciplinary Hall, R.A., Carter, G.S., 2011. Internal tides in Monterey Submarine Canyon. Journal
of Physical Oceanography 41, 186–204.
discussions and investigations. Harris, P.T., Barrie, J.V., Conway, K.W., Greene, H.G., this issue. Hanging canyons of
Haida Gwaii, British Columbia, Canada: fault-control on submarine canyon
geomorphology along active continental margins. Deep-Sea Research II. this
issue [http://dx.doi.org/10.1016/j.dsr2.2013.09.010].
Acknowledgments Harris, P.T., Whiteway, T., 2011. Global distribution of large submarine canyons:
geomorphic differences between active and passive continental margins.
The authors would like to thank NOC Southampton, and Marine Geology 285, 69–86.
Ingels, J., Tchesunov, A.V., Vanreusel, A., 2011. Meiofauna in the Gollum Channels
especially IFREMER for supporting the first INCISE meeting and and the Whittard Canyon, Celtic Margin – how local environmental conditions
for help in starting up the network. We are also very grateful shape Nematode structure and function. PLoS One 6, e20094.
towards the INCISE committee (Rob Hall, Peter Harris, Aaron Juniper, S.K., Matabos, M., Mihaly, S., Ajayamohan, R.S., Gervais, F., Bui, A.O.V., 2013.
A year in Barkley Canyon: a time-series observatory study of mid-slope benthos
Micallef, Joshu Mountjoy and Nathalie Valette-Silver) for the
and habitat dynamics using the NEPTUNE Canada network. Deep-Sea Research
fruitful discussions and support. V. Huvenne is funded by the Part II: Topical Studies in Oceanography 92, 114–123.
Natural Environment Research Council MAREMAP programme, Kenchington, E., Cogswell, A., MacIsaac, K., Beazley, L., Law, B., Kenchington, T., this
and through a Starting Grant of the European Research Council issue. Limited depth zonation among bathyal epibenthic megafauna of the
Gully submarine canyon, northwest Atlantic. Deep-Sea Research II. [http://dx.
(ERC project CODEMAP, Grant no 258482). doi.org/10.1016/j.dsr2.xxxx.xx.xxx].
Lastras, G., Canals, M., Urgeles, R., Amblas, D., Ivanov, M., Droz, L., Dennielou, B.,
Fabres, J., Schoolmeester, T., Akhmetzhanov, A., Orange, D., Garcia-Garcia, A.,
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