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A Tale of Two Basins An Integrated Physical and Biological Perspective of The Deep Arctic Ocean 2015 Progress in Oceanography
A Tale of Two Basins An Integrated Physical and Biological Perspective of The Deep Arctic Ocean 2015 Progress in Oceanography
A Tale of Two Basins An Integrated Physical and Biological Perspective of The Deep Arctic Ocean 2015 Progress in Oceanography
Progress in Oceanography
journal homepage: www.elsevier.com/locate/pocean
UiT - The Arctic University of Troms, Department of Arctic and Marine Biology, 9037 Troms, Norway
School of Fisheries and Ocean Sciences, Institute of Marine Science, University of Alaska Fairbanks, 905 Koyukuk Drive, Fairbanks, AK 99775-7220, USA
c
P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Nakhimovsky Prospect 36, Moscow 117218, Russia
d
Fisheries and Oceans Canada, Institute of Ocean Sciences, 9860 West Saanich Road, Sidney, British Columbia V8L 4B2, Canada
b
a r t i c l e
i n f o
Article history:
Available online 8 August 2015
a b s t r a c t
This review paper integrates the current knowledge, based on available literature, on the physical and
biological conditions of the Amerasian and Eurasian basins (AB, EB) of the deep Arctic Ocean (AO) in a
comparative fashion. The present day (Holocene) AO is a mediterranean sea that is roughly half continental shelf and half basin and ridge complex. Even more recently it is roughly two thirds seasonally and one
third perennially ice-covered, thus now exposing a portion of basin waters to sunlight and wind. Basin
boundaries and submarine ridges steer circulation pathways in overlying waters and limit free exchange
in deeper waters. The AO is made integral to the global ocean by the Northern Hemisphere Thermohaline
Circulation (NHTC) which drives Pacic-origin water (PW) through Bering Strait into the Canada Basin,
and counter-owing Atlantic-origin water (AW) through Fram Strait and across the Barents Sea into
the Nansen Basin. As a framework for biogeography within the AO, four basic, large-scale circulation systems (with L > 1000 km) are noted; these are: (1) the large scale wind-driven circulation which forces the
cyclonic Trans-Polar Drift from Siberia to the Fram Strait and the anticyclonic Beaufort Gyre in the southern Canada Basin; (2) the circulation of waters that comprise the halocline complex, composed largely of
waters of Pacic and Atlantic origin that are modied during passage over the Bering/Chukchi and
Barents/Siberian shelves, respectively; (3) the topographically-trapped Arctic Circumpolar Boundary
Current (ACBC) which carries AW cyclonically around the boundaries of the entire suite of basins, and
(4) the very slow exchange of Arctic Ocean Deep Waters. Within the basin domain two basic water mass
assemblies are observed, the difference between them being the absence or presence of PW sandwiched
between Arctic Surface Waters (ASW) above and the AW complex below; the boundary between these
domains is the Atlantic/Pacic halocline front. Both domains have vertical stratication that constrains
the transfer of nutrients to the surface layer (euphotic zone), thus leading to their oligotrophic state, particularly in the more strongly stratied Pacic Arctic where, despite high nutrient values in the inow,
convective reset of surface layer nutrients by haline convection in winter is virtually absent. First and
multi-year sea ice drastically alters albedo and insulates the underlying water column from extreme winter heat loss while its mechanical properties (thickness, concentration, roughness, etc.) greatly affect the
efciency of momentum transfer from the wind to the underlying water. Biologically, sea ice algal growth
in the basins is proportionally almost equal to or exceeding phytoplankton production, and is a habitat
and transport platform for sympagic (ice-associated) fauna. Owing to nutrient limitation due to strong
stratication and light limitation due to snow and ice cover and extreme sun angle, primary production
in the two basin domains is very low compared to the adjacent shelves. Severe nutrient limitation and
complete euphotic zone drawdown in the AB favors small phytoplankton, a ubiquitous deep chlorophyll
maximum layer, a low f-ratio of new to recycled carbon xation, and a low energy food web. In contrast,
nutrients persist albeit in low levels in the western EB, even in summer, suggesting light limitation,
heavy grazing or both. The higher stocks of nutrients in the EB are more conducive to marginal ice blooms
than in the AB. The large-scale ocean currents (NHTC and ACBC) import substantial expatriate, not locally
reproducing zooplankton biomass especially from the adjoining subarctic Atlantic (primarily Calanus nmarchicus), but also from the Pacic (e.g., Pseudocalanus spp., Neocalanus spp. and Metridia pacica). These
advective inputs serve both as source of food to resident pelagic and benthic biota within the basins, and
Corresponding author at: UiT - The Arctic University of Troms, Department of Arctic and Marine Biology, 9037 Troms, Norway. Tel.: +47 776 44382.
E-mail address: bodil.bluhm@uit.no (B.A. Bluhm).
http://dx.doi.org/10.1016/j.pocean.2015.07.011
0079-6611/ 2015 Elsevier Ltd. All rights reserved.
90
Nomenclature
Acronyms
AB
Amerasian Basin
AO
Arctic Ocean
AOI
Arctic Oscillation Index
ACBC
Arctic Circumpolar Boundary Current
ASW
Arctic Surface Waters
AW
BSB
EB
FSB
NCP
PW
Atlantic-origin Water
Barents Sea Branch
Eurasian Basin
Fram Strait Branch
net community production
Pacic-origin water
as potential grazers exerting top down control on limited phytoplankton resources. Benthic organisms
within the AO basin show previously unappreciated biodiversity and surprising dispersion of species
given the isolation of individual basins and low vertical carbon ux and resulting biomass. Larval dispersion is aided by the large-scale ows and perhaps, we hypothesize in the deep benthos by convective
updrafts driven by geothermal heating. Zooplankton diversity, in contrast, is low, but again faunal assemblages are equally distributed between the EB and AB. Species pools of both pelagic and benthic communities change more with water depth rather than laterally, with the exception of expatriates and rare
species, with close ties to todays North Atlantic biogeographic region. Climate related change in the
AO is thus manifest at signicantly differing time scales. Throughout 90% of the Pleistocene the AO
has existed in glacial mode, with narrow continental shelves, greatly restricted river inow, thicker and
perhaps immobile sea ice, and total blockage of exchange with the Pacic Ocean. During the Holocene,
on shorter time scales of 1000100 years, signicant changes in high latitude climate are tied to changes
in temperature and perhaps moisture delivery patterns. The Arctic also experiences signicant
multi-decadal variability; however, the pace of change over the past three decades has been without
precedent. Within the basin interior the ice is now thinner and less compact, and thus more responsive
to wind stress (forcing and mixing). Concurrent with sea ice melt and increased river ow, the accumulation of fresh water and the stratication have increased, thus constraining vertical nutrient ux affecting phytoplankton size distributions, limiting primary production in parts of the basins now and likely in
the future, and increasing vulnerability to acidication. In addition, sea ice is now retreating on an annual
basis past the shelf break, exposing basin waters directly to sunlight and wind forcing. Thus, upwelling
favorable winds (generally from east to west) can now directly and efciently drive shelf-break upwelling, and draw nutrients from subsurface basin waters onto the shelf; at the same time upwelling favorable winds will also create onshore pressure gradients over the slope and basin which will act to slow or
block the ow of waters in the ACBC, and thus alter advective pathways of both abiotic and biotic materials. Given the opening of a new ocean to multiple user groups, we expect that the central AO will play an
increasing larger role both in the research and political arenas in the future, and we encourage pan-Arctic
international collaboration over focus on territorial boundaries.
2015 Elsevier Ltd. All rights reserved.
1. Introduction
Following earlier descriptions we here refer to the panarctic
domain north of Bering Strait in the Pacic and
Greenland-Scotland Ridge in the Atlantic as the Arctic Mediterranean; the Norwegian, Iceland and Greenland seas south of Fram
Strait and west of the Barents Sea comprise the Nordic Seas, and
the remainder, including both shelf and basin domains, is the Arctic Ocean (AO). We refer to the central basins at the Amerasian
Basin (AB) and the Eurasian Basin (EB). Our view of this ocean
has changed over time. It has changed because we have increased
our understanding of the area through painstaking scientic measurements since the time of the early Arctic explorers, to todays
increasingly diverse suite of measurements by icebreakers, ice
camps, moorings, ice-tethered prolers and satellites (Fig. 1). It
has also changed because the system itself is in a rapid state of
transition (ACIA, 2005; Wassmann and Lenton, 2012; Polyakov
et al., 2013; Bhatt et al., 2014). And central to understanding the
AO system now and into the future is to understand its least studied component: the deep basins.
At the onset of his historic voyage of 18931896 Nansen
believed the entire AO to be a shallow sea (Nansen, 1902). And
while Nansen discovered that a deep basin occupied the central
AO, its trans-arctic ridge systems were unsuspected until the middle of the 20th century (cf. Fig. 2a) which initially were deduced
from tide and temperature data (Fjeldstad, 1936; Worthington,
1953). Modern charts (Fig. 2b) now reveal two main basins, the
Eurasian (EB) and Amerasian basins (AB), separated by the Lomonosov Ridge. The EB is further separated into the Nansen and
Amundsen basins by the Gakkel Ridge, and the AB into the
Makarov and Canada Basin by the AlphaMendeleyev Ridge.
Though the job of accurately mapping the AO bathymetry is far
from complete, we recognize the important fact that more than
half (53%) the area of the present day AO is continental shelf, and
the remainder is the slope/rise/basin/ridge complex; (Fig. 2c;
Aagaard et al., 1985; Jakobsson et al., 2012). To simplify the complicated bathymetry of the Arctic Mediterranean, and to link physical and morphometric properties with biological distributions and
functions, we follow Carmack and Wassmann (2006) and divide
the Arctic Mediterranean into fundamental hydro-morphological
domains. These are the inow shelves, the interior shelves, the
outow shelves, the Riverine Coastal Domain (Carmack et al.,
2015), the two main basins, and the ridge and borderland features
(cf. Fig. 2d). This distinction is critical to the understanding of basin
dynamics and biogeography owing to the importance of
shelf-basin interactions and inter-domain advection.
Fig. 1. Bar graph illustrating the number of ice camp and ship-based expeditions
taking place in the basins of the Arctic Oceans over the past six decades.
91
The large ratio of shelf to basin area is one of the truly unique
features of the modern AO; it is, however, a signature that has
changed often and rapidly in geologic time. The Arctic Basins began
to develop with the AB rst ca. 140150 Myrs ago, and then the EB
ca. 5070 Myrs ago (cf. Backman and Moran, 2009). The AOs deep
connection to the Pacic closed ca. 80 Myrs ago (Marincovich et al.,
1990), leaving Fram Strait (sill depth 2600 m) as its only deep connection to the global ocean. The AO became the modern Arctic ca.
4 Myrs ago with the closure of the Central American Seaway
(which began to freshen the Pacic relative to the Atlantic)
(Zauker et al., 1994; Haug et al., 2001; Hasumi, 2002) and the
opening Beringia Seaway (which allows the fresh Pacic water to
ow back into the Atlantic via the AO pathways) (Coachman and
Barnes, 1961; Carmack and McLaughlin, 2011). Ice core records
show that during this time, the Earths climate system began to
resonate with orbital cycles, leading perhaps to the
glacial-interglacial periods extant today (cf. Fedorov et al., 2006).
In glacial mode the AO is almost all deep basin (except for the Barents Sea) and is fed only by inow from the Atlantic. In contrast,
Fig. 2. Evolution of understanding of the Arctic basins bathymetry and hydro-morphometry: (a) bathymetric map of the Arctic Ocean at the mid-20th Century (Shirshov,
1944); (b) most-recent IBCAO map of the Arctic Ocean (Jakobsson et al., 2012); (c) simplied bathymetry and place names used in this paper; and (d) highly idealized
typology of the Arctic Mediterranean based on its hydro-morphological domains: AW is Atlantic Water; PW is Pacic Water; RCD is Riverine Coastal Domain (see Carmack
et al., 2015); AO is Arctic Outows; and arrows denote component ow directions.
92
93
Fig. 3. Historical perspective on Arctic Ocean connectivity and modern ice cover changes: (a) map from the mid-19th century illustrating the then prevailing notion of a
central Arctic Ocean kept ice-free by inowing ocean currents (from Overland et al., 2011); (b) highly idealized schematic of ows linking arctic and subtropical waters
according to the requirement to materials from regions where there is excess to regions where there is decit (from Sandstrm, 1919); (c) map showing present day estimates
of transports and average T and S properties of the inows and outows based on an inverse model applied to boundary uxes in summer 2005 (Tsubouchi et al., 2012), and
mass balances estimates at Fram Strait (2.0 2.7 Sv, 19972006, Schauer et al., 2008), at the Barents Sea Opening (2.0 Sv, 19972007, Smedsrud et al., 2010), at Bering Strait
(0.8 0.2 Sv, 19912004, Woodgate et al., 2005; Melling et al., 2008) and Davis Strait (2.3 0.7 Sv for 20042005; Curry et al., 2011); and (d) map comparing the Arctic
Ocean ice cover (September minimum) for the 19792000 mean versus 2012 and overlaid on ocean bathymetry.
where such water abounds to where it is rare . . . the complimentary physical processes whereby water is added to or drawn from
a certain layer adapt themselves so as to balance one another in
rate (Fig. 3b). While the expeditions of the late 1800s and early
1900s, for example Nansens Fram expedition in 18931896
(Nansen, 1902), provided proof against the open Polar Sea concept,
they did result in early evidence of Atlantic inows to the deep
basins and of the primary direction of ice drift across the Arctic
Basin via Transpolar Drift (Rudels, 2012). These ndings were later
to be conrmed by observations from Russian ice drift stations
(called North Pole) that began in 1937 (Shirshov and Fedorov,
1938). The associated uxes of mass, heat and salt by Atlantic
and Pacic waters have recently been summarized under the banner of the Arctic and Subarctic Ocean Fluxes program (Dickson
et al., 2007, 2008; Fig. 3c). While the notion of an ice free polar
94
Fig. 4. Horizontal maps of annual mean near-surface (20 m) (a) salinity and (b) density north of 20N for the Northern Hemisphere illustrate the basic estuarine circulation
forced by low salinity and low density waters entering from the Pacic and more saline and denser waters entering from the Atlantic. Data source: http://odv.awi.de/en/data/
ocean/world_ocean_atlas_2009/.
(mainly Nares Strait) are exported into the convective gyres of the
subarctic North Atlantic and then join the global thermohaline circulation which carries low-salinity waters back to the low latitudes, and therefore contributes to closing the global freshwater
loop (cf. Carmack and McLaughlin, 2011; for summary).
For the purpose of understanding biogeographical distributions
within the AO, and to serve as a backdrop for an ecology of advection (c.f. Wassmann et al., 2015), it is useful to recognize four basic
large-scale (with L > 1000 km) circulation systems; these are: (1)
the large scale wind-driven circulation of ice and the upper ocean
(0 m to 50 or 150 m, depending on location) which forces the
cyclonic Trans-Polar Drift from Siberia to the Fram Strait and the
anticyclonic Beaufort Gyre in the southern Canada Basin
(Fig. 5a); (2) the circulation of waters that comprise the halocline
complex, composed largely of waters of Pacic and Atlantic origin
that are modied by freeze/thaw processes during passage over
the Bering/Chukchi and Barents/Siberian shelves, respectively
(Jones and Anderson, 1986; Aksenov et al., 2011); (3) the
topographically-trapped Arctic Circumpolar Boundary Current
(ACBC) which carries AW cyclonically around the boundaries of
the entire suite of basins (Aagaard, 1989; Aksenov et al., 2011;
Rudels et al., 2013; Fig. 5c); and (4) the very slow exchange of
AO deep waters which form initially in the Greenland Sea, enter
through Fram Strait, and spread within the basin interior sequentially to the Nansen, Amundsen, Makarov and Canada basins
(Macdonald et al., 1993; Schlosser et al., 1997; Rudels et al.,
2012; Fig. 5d). Exchange of the latter from basin to basin is laterally
constrained by the sequence of sills, but is aided in vertical motion
by geothermal heating (Timmermans et al., 2003; Carmack et al.,
2012). A summary of mid-water sources, pathways and associated
fronts are shown in Fig. 5d. Details of this circulation typology are
discussed next.
Surface Circulation: Much of what we know of surface circulation patterns stems from early observation collected from Russian
ice stations (cf. Coachman and Aagard, 1974; Frolov et al., 2005)
and was deduced from observations of sea ice drift, starting with
Nansen (1902) and culminating in the International Arctic Buoy
Program (IABP) (Rigor et al., 2002; Prman et al., 2004). Sea ice
and water masses lying above the cyclonically spreading Atlantic
waters (including the ACBC) are primarily wind-driven. Two basic
systems are manifest: (1) the cyclonic (Atlantic) Arctic bounded to
the east by the Siberian shelves and to the west by the Trans-Polar
95
Fig. 5. Schematic representations of Arctic Ocean circulation: (a) Surface circulation of the Arctic Ocean as shown by dynamic topography (20/400 dbar) (World Ocean
Database 2013, (b) summary of mid-water halocline sources, ows and associated fronts (blue shows Pacic-origin waters, maroon shows Atlantic-origin waters, thick
maroon line depicts the front between them) (after McLaughlin et al., 1996); (c) schematic representation of the Arctic Circumpolar Boundary Current system (ACBC) derived
from Atlantic water inows (after Aksenov et al., 2011; Rudels et al., 2013); and (d) schematic representation of deep water exchange (Aagaard et al., 1985). BG is the Beaufort
Gyre, BSB is the Barents Sea Branch, FSB is the Fram Strait Branch, GG is the Greenland Gyre, NAC is the Norwegian-Atlantic Current, NCC is the Norwegian Coastal Current,
TPD is the Transpolar Drift.
Drift and (2) the anticyclonic Beaufort Gyre over the Canada Basin
(Fig. 5a). The cyclonic ow over the EB and some of the Makarov
Basin denes a fundamentally divergent gyre, while the anticyclonic ow of the Beaufort Gyre denes a convergent system
(Fig. 5a); an important distinction in terms of regional stratication
and nutrient availability to the euphotic zone.
In recent years our understanding has been advanced by the use
of satellite observations (sea surface height and bottom pressure)
and much expanded in situ observations by ships, submarines, aircraft, ice camps, moorings and ice-tethered prolers (Morison
et al., 2012). From this has come a much deeper appreciation of
the degree of variance from long-term mean in both velocity and
in the spreading pathways of water masses. Proshutinsky and
Johnson (1997) used a modeling approach to demonstrate two
modes of AO circulation according to patterns in atmospheric forcings. It was later suggested that such variability could inuence the
storage and release of freshwater within the Beaufort Gyre into the
convective gyres of the Nordic Seas downstream (Proshutinsky
et al., 2002). Subsequently, a regional-scale time series carried
out in the Canada Basin since 2003 has revealed a trend and large
year-to-year variability in freshwater storage and strength of surface circulation within the Beaufort Gyre (McLaughlin et al.,
2009; Proshutinsky et al., 2009; Krisheld et al., 2014).
Kwok et al. (2013) applied satellite data collected over a 28-year
period (19822009) to summarize the variance and shifts in decadal drift patterns in ice drift velocity and compared these to geostrophic ow elds using the polarity of the Arctic Oscillation as a
backdrop for atmospheric forcing. Mean circulation speeds over
the basin are of order 24 cm/s, with stronger ows along the
southern edge of the Beaufort Gyre, in the Transpolar Drift, and
especially in the Fram Strait Outow. These authors further report
a net strengthening of the Beaufort Gyre and the Transpolar Drift,
especially during the last decade, with over 90% of the AO displaying a positive trend in drift speed, and a decline in multiyear sea ice
coverage. Importantly, they note the spatially averaged trends
from 20012009 in drift speeds (winter +24%/decade, summer
+18%/decade) are not explained by the much smaller trends in
wind speeds (winter +1.5%/decade, summer: 3.4%/decade),
96
with positive trends in drift speed in regions with reduced multiyear sea ice coverage. The increased responsiveness of ice drift to
geostrophic wind is consistent with a thinner and weaker seasonal
ice cover and suggests large-scale changes in the air-ice-ocean
momentum balance related to reductions in ice strength and
concentration.
Halocline waters: Formation and subsequent circulation of halocline waters within the AO basins remains poorly known, and is
likely due to a combination of mechanisms. Aagaard et al. (1981)
and Melling and Lewis (1982) discussed the importance of sea
ice formation and brine drainage in modifying shelf-derived
waters, and Killworth and Smith (1984) demonstrated the importance of Pacic inows to halocline formation. Rudels et al.
(1996) proposed that wintertime convection in the Nansen Basin
followed by summertime capping by ice melt and outow of fresh
shelf water contributed to halocline formation. Kikuchi et al.
(2004) further discussed the formation of convectively-formed
waters in the EB and its spatial and temporal variability, noting
that large-scale advance and retreat into the Amundsen Basin
appeared to coincide with ocean circulation and frontal shifts associated with increased cyclonic circulation in the atmosphere. Jones
and Anderson (1986) used chemical distributions to distinguish
between what they termed the upper (Pacic-derived) and lower
(Atlantic-derived) halocline components. Bauch et al. (2014) use
hydrochemical and oxygen isotope data do document an
along-slope front between shelf, slope and central EB waters in
the Laptev Sea, and conclude that halocline waters above the slope
are derived from upstream advection. McLaughlin et al. (1996)
argued that a front, the Atlantic-Pacic halocline front, spans the
Arctic basins where the Atlantic-derived waters subduct below
the Pacic-derived waters (Fig. 5b, also Karcher et al., 2012). They
further suggest that this front roughly aligns with and shifts
between the Lomonosov and AlphaMendeleev Ridge on decadal
time scales.
The inow of PW is likewise complicated, with branches developing south of Bering Strait from Anadyr Coastal Current waters,
central Bering Shelf waters, and the Alaska Coastal Current waters,
and continuing north across the Chukchi Sea via Herald, Hanna and
Barrow canyons (Weingartner et al., 1998, 2013; Pickart et al.,
2005, 2010). Due to the considerable widths of the Bering and
Chukchi shelves, and thus the long crossing time, these inows
are modied en route to become the denser, winter and lighter,
summer varieties of PW, with each branch undergoing differing
geochemical transformations (Nishino et al., 2013). Upon entry
into the AB, the tendency for PW to circulate in the cyclonic sense
of the ACBC is opposed by the anticyclonic, wind-driven Beaufort
Gyre (cf. Shimada et al., 2006). Using modeling results, Aksenov
et al. (2011) argued that cyclonic ow along the Alaska shelf and
upper slope in the AB is forced by steric sea level differences
between the Pacic and Arctic. The conuence of Pacic waters
with halocline source waters draining from the Siberian shelves
results in the Atlantic/Pacic Halocline front, separating the two
domains (Fig. 5c).
The Arctic Circumpolar Boundary Current: The input of AW to the
AO has been studied for over a century but, because of its complexity and variability, pathways and uxes are still subject to huge
uncertainties; a recent census of uxes through the main gateways
is given by Beszczynska-Mller et al. (2011). Between 8 and 9 Sv
(1 Sv = 106 m3 s1) enter the Nordic Seas over the Greenland Scotland Ridge (sill depth 800 m) and of this roughly half continues
on to the AO. Of the AW continuing north, about half enters the
AO via Fram Strait as the Fram Strait Branch (FSB) and subducts
below Arctic Surface waters (ASW) north of Svalbard. The other
half rst crosses the Barents and westernmost Kara seas, subducts
along the Atlantic Polar Front, continues across the eastern Barents
Sea and then enters the St. Anna Trough (Rudels et al., 2013). Here,
97
More rapid ows are expected along basin and ridge slopes, and
through narrow gaps in the ridges. For example, the Lomonosov
Ridge separates the EB and AB with average depths below sea level
between 1000 and 1400 m, but with a narrow gap of sill depth
1870 m near 88200 N, 148E. (Bjrk et al., 2007). This gap leads
from the Makarov Basin side of Intra Basin, a sub-basin atop the
Lomonosov Ridge. Timmermans et al. (2005) apply hydraulic theory to estimate a ow over the Lomonosov Ridge of
0.25 106 m3 s1, but Timmermans and Garrett (2006) speculate
that ows of this magnitude are not reaching fully to the bottom.
Bjrk et al. (2007), however, present hydrographic data giving evidence that ow is actually from the Makarov to the Amundsen
Basin, and subsequently interowing at intermediate depths
(17002000 m) as a return ow from the Canadian Basin to the
EB and the Nordic Seas. Note that this direction of the water overow is opposite to that previously proposed by Jones et al. (1995)
and Timmermans et al. (2005) and discussed as intermittent ow
by de Middag et al. (2009). Geologic observations (Bjrk et al.,
2007) and modeling results (Isachsen et al., 2003) are consistent
with strong current activity at the ridge crest, thus aiding planktonic dispersion.
Vertical motions may also play an important role in dispersion
of biota. For example, an outstanding feature of the AO deep water
is its near homogeneity over thicknesses exceeding 1200 m within
the Canada Basin and 800 in the Amundsen Basin, and capped by
a temperature minimum (Timmermans et al., 2003; Bjrk and
Winsor, 2006; Carmack et al., 2012). In the Canada Basin, for example, the deep waters vary in potential temperature (h) by less than
0.001 C between 2700 m and the bottom, a feature that
Timmermans et al. (2003) and Carmack et al. (2012) ascribe to
geothermal heating and vertical convection. Salinity in the Canada
Basin increases with depth to 2700 m, but, like h, is nearly constant below this depth, at S = 34.957 psu. Bjrk and Winsor
(2006) reported similar, near-homogenous deep waters in the EB,
which they also attributed to geothermal heating. Using a
time-series of deep water properties, Carmack et al. (2012) found
that Canada Basin deep waters below 2700 m warmed at a rate
of 0.0004 C yr1 between 1993 and 2010. This rate is slightly less
than that to be expected from the reported geothermal heat ux
(50 mW m2; cf. Langseth et al., 1990). Using this heat ux they
estimated a vertical velocity scale of of 0.8 mm s1, and thus for
the 1000 m-thick Canada Basin Deep Water the time-scale for convection is calculated to be 15 days. This value suggests rapid vertical mixing of the deep waters, particularly in relation to its long
isolation age. The deep waters found above the lower continental
margin of the deep basin maintain higher temperatures than those
in the basin interior, consistent with geothermal heat distributed
through a shallower water column.
Lateral, inter-basin exchange of water along isopycnal surfaces
at or near sill depths may effectively disperse biota. Carmack
Table 1
Comparison between water mass properties of and river inow into the Eurasian and Amerasian Basins. Halocline values are typical of summer conditions.
Eurasian basin
Amerasian basin
Reference
1.7 to 4 C/33.834.8
One halocline/dS = 2
13 C/34.834.9
0.9 C/34.92
250
1015
Ob, Lena, Yenisey, Pechora,
Kolyma, Severnaya Dvina
65%
17%
58
1.7 to 4 C/28.034.4
Stepwise/dS up to 10
0.60.8 C/34.8034.85
0.5 C/34.95
450
15
Mackenzie, Yukon (indirect)
JOIS
JOIS
McLaughlin et al. (2009)
Bjrk and Winsor (2006) and Carmack et al. (2012)
Macdonald et al. (1993) and Schlosser et al. (1997)
Codispoti et al. (2013)
Holmes et al. (2002)
21%
73%
1
98
Fig. 6. Representative proles of (a) potential temperature (h), (b) salinity (S) and (c) h/S correlations for the four sub-basins of the Arctic Ocean: green is Nansen Basin, red is
Amundsen Basin; yellow is Makarov Basin and blue is Canada Basin. Water mass assemblies are distinguished by the absence of presence of Pacic Waters sandwiched within
the halocline. Note that the depth axis is plotted as square root of depth to better dene the upper ocean where detail is greatest. Data from the JOIS program 200504 and
from Polarstern ARKXII-1.
Surface waters of the AO are comprised of a base of either Atlantic or Pacic origin waters (depending on basin location) and
diluted by river inputs, ice melt and net precipitation
(Yamamoto-Kawai et al., 2009). Far more river water is supplied
to the EB than to the AB (Holmes et al., 2002); however, the residence time of these waters is relatively short, of order two years,
as it is carried quickly from the basin into Fram Strait by the cyclonic ow of the EB and the Trans-Polar Drift (Anderson et al., 1989).
In contrast, the accumulation of river water is especially pronounced in the Canada Basin, where the convergent winds of the
atmospheric Beaufort High accumulate low salinity waters of both
North American and Siberia within the anticyclonic Beaufort Gyre,
making this gyre the most strongly stratied component of the AO
(Aagaard and Carmack, 1989; Proshutinsky et al., 2009). The resulting heterogeneity in the distribution of freshwater components is
reected in maps of surface salinity (Fig. 4a).
Pacic Waters enter from the Bering Sea through Bering Strait,
transit the Chukchi Sea and enter the deep basins along three main
branches: one associated with Barrow Canyon; one east of Hannah
Shoals; and one following Herald Canyon (Weingartner et al., 1998,
2005; Aagaard et al., 2006). Inowing waters include nutrient-rich
water from the Gulf of Anadyr and fresher, lower nutrient Alaskan
Coastal Cater (Walsh et al., 1989). While crossing the wide Chukchi
Shelf, is modied seasonally by biological production, heat
exchange, ice formation and melting, and interaction with
99
Fig. 7. Section of salinity (S) colored by potential temperature (h) for an XCTD section crossing the Amerasian Basin; inset shows station locations. Note that the Atlantic/
Pacic Halocline Front is evident in the steeply-sloping isohalines in the Canada Basin leading up to the Alpha-Mendeleev Ridge, and by the warmer Pacic varieties of
halocline waters extant within the Canada Basin (from Kikuchi, Itoh, Eert and Williams, pers. comm.). Black vertical lines in bottom gure indicate locations of XCTD
deployments.
tantly, with regards to this paper, is the fact that ice retreat is
now exposing surface waters over the deep basins; in 2012
approximately 40% of basin area (with depth >400 m) was ice free.
While not yet quantied, the effects of increased wind and solar
forcing on the upper layers of the deep basins are likely to be
substantial.
Wind and atmospheric thermodynamic forcing were the main
causes for the record Arctic sea ice retreat in summer2007 (e.g.,
Perovich et al., 2008). However, AW heat was likely important
for preconditioning Arctic sea ice by making it thinner over several
preceding decades and thus contributing to the extreme retreat
(Polyakov et al., 2010). A similar situation exists for incoming
warm Pacic Summer Water spreading off shelf and into the Beaufort Gyre in the Canada Basin (Shimada et al., 2006). Another feature of observed decline in sea ice extent is that the trend is not
strictly linear, but instead consists of a series of punctuated
changes, e.g. in 1989, 1998 and 2007 (see Perovich et al., 2014
for time series). One plausible explanation is that ice thickness
and strength is slowly and inexorably decreased, year by year, by
internal forcing related to increased heat advection by the atmosphere and ocean, and then shocked into a new dynamical state
by external forcings in extreme years. From the perspective of
complex systems behavior, thermodynamic forcing by the atmosphere and ocean acts as the slow variable to reduce ice cover
strength and resilience to external forcing, so that when exceptional wind patterns do occur, such as in 2007 (Stroeve et al.,
2012; Arctic Council, 2013), ice is readily exported south to melt
in the Nordic Seas.
The above noted changes in sea ice are biologically signicant
since sea ice and snow play several key physical and biogeochemical roles in the Arctic marine system. From a heat budget perspective, the presence of sea ice drastically alters albedo and insulates
the underlying water column from extreme winter heat loss. The
mechanical properties of sea ice (thickness, concentration, roughness, etc.) greatly affect the efciency of momentum transfer from
100
Fig. 8. Horizontal maps showing distributions of nutrient concentrations: (a) surface nitrate; (b) nitrate at 200 m; (c) surface phosphate; (d) phosphate at 200 m; (e) surface
silicate; and (f) silicate at 200 m. Data derived from the Hydrochemical Atlas, CARINA, Codispoti et al., 2013 and JOIS (http://catalog.data.gov/dataset/hydrochemical-atlas-ofthe-arctic-ocean-nodc-accession-0044630, http://cdiac3.ornl.gov/waves/discrete/, http://www.nodc.noaa.gov/archive/arc0034/0072133/1.1/data/0-data/).
the wind to the underlying water, with a thinner and loosely consolidated ice cover being more effective in driving ocean currents.
Freezing of sea ice within the basins and subsequent local melting
is observed to increase stratication, while the export and melting
of ice in the adjacent North Atlantic is a signicant component in
the AOs freshwater budget (Aagaard and Carmack, 1989). Sea ice
melt water also has differing geochemical properties from ASW,
such as heavier d18O values which can be used as a tracer
(Macdonald et al., 1999), and lower alkalinity, a property that
may exacerbate ocean acidication (Yamamoto-Kawai et al.,
2009). Biologically, sea ice is the site of ice algal growth and transport, a habitat for ice meiofauna, zooplankton and small sh, and a
platform on which marine mammals may travel, hunt and breed
(Bluhm and Gradinger, 2008).
Two aspects of sea ice reduction are worth special attention
with regard to the deep basins. First, prior to the 21st century
the sea ice margin seldom retreated beyond the shelf-break, and
thus basin waters were seldom exposed to sunlight and wind. In
2012, in contrast, roughly 40% of the area of the deep basins was
exposed (Fig. 2d). This would increase solar radiation to basin
waters, enhance wind mixing and increase shelf-basin exchange
(see Section 6.3). The full physical and ecological consequences
of this new normal state (Wood et al., 2013; Jeffries et al., 2013)
are currently a matter of debate, and on a regional basis, primary
production may either increase (Arrigo et al., 2008, for
light-limited shelf areas) or decrease (McLaughlin and Carmack,
2010, for nutrient impoverished basins; Section 7). A second
impact is that the reduction of ice in summer now allows enhanced
gas exchange with the atmosphere, notably CO2, and this has been
implicated as a factor in increasing the acidity of AO surface waters
over the basins (Yamamoto-Kawai et al., 2011).
3.4. River inow and other freshwater in the basins
Of all global oceans the AO is the most riverine, covering only
3% of the global ocean surface area but capturing 10% of global
river runoff within its mediterranean boundaries (Carmack,
2000). Further, river inputs are increasing with global warming
(Peterson et al., 2002, 2006), albeit with disproportional increase
in Eurasian river run-off (McClelland et al., 2006). River inputs to
the AO also play a role in biogeochemical processes. Rivers bring
turbidity that can counteract enhancing effects of nutrients by
blocking sunlight to primary producers and clogging
lter-feeders (Syvitski et al., 1989). Direct nutrient inputs from rivers are relatively small compared to the advective inputs from the
Atlantic and Pacic Oceans (Codispoti and Owens, 1975), and likely
support only 10% of total NCP (Gordeev et al., 1996; Gordeev,
2000; see also Section 4.1). Alkalinity values, however, are typically
lower in incoming rivers than in AO waters, and their mixing with
and inclusion in surface waters acts in increase local vulnerability
to ocean acidication (Yamamoto-Kawai et al., 2011).
Liquid fresh water is also supplied to the AO through inputs
from sea ice melt and net precipitation and, relative to an appropriate reference salinity by the Norwegian Coastal Current and
inows from the Pacic Ocean through Bering Strait. Sinks of liquid
fresh water include export through the Canadian Arctic Archipelago and the western Fram Strait, and export of sea ice. Each of
the sources contributes uniquely to halocline formation and structure within the system, and to a variety of circulation and mixing
processes that affect biological distributions.
An Arctic freshwater budget for sources, sinks and storage was
rst produced by Aagaard and Carmack (1989), and has been subsequently updated by Serreze et al. (2006), Dickson et al. (2007),
Tsubouchi et al. (2012) and Haine et al. (2015). Most estimates of
freshwater content are based on the choice of reference salinity
(for discussion, see Carmack et al., 2008). Aagaard and Carmack
(1989) and Serreze et al. (2006) used 34.8, the mean salinity of
the AO, while Dickson et al. (2007) calculated content relative to
a salinity of 35, the salinity of incoming AW. Using available historical data, Serreze et al. (2006) calculated a net AO freshwater
export of 9200 km3 yr1 and an import of 8500 km3 yr1, leaving
a net imbalance of 700 km3 yr1. Using a similar approach, but
with a different reference salinity and study area, Dickson et al.
(2007) computed an export of about 9500 km3 yr1. Tsubouchi
et al. (2012) applied an inverse model to constrain ux estimates
for volume, heat and freshwater around the AO boundary (waters
101
above 1000 m) for a 32-day period in summer 2005. They calculated mean properties for water entering the Arctic to be
h = 4.49 C and S = 34.50, and for water leaving the Arctic, including
sea ice, h = 0.25 C and S = 33.81 (Fig. 3c). They calculated a corresponding volume transport into the Arctic of 9.2 Sv, an export of
9.3 Sv, and a net oceanic and sea ice freshwater ux of
187 48 mSv (Fig. 3c).
Regional circulation processes play a huge role in the distribution and pathways of freshwater components within the basins.
Initial river inow at source creates a coastal-trapped Riverine
Coastal Domain owing from west to east and rimming much of
the inner continental shelf (cf. Carmack and McLaughlin, 2000;
Carmack et al., 2015, Fig. 2d). Upon exiting the shelves, for example
during upwelling favorable wind events, river water mixtures
serve to freshen the surface waters within the basins. In general,
wind-driven currents serve either to store fresh water, as is the
case for the convergent Beaufort Gyre, or to export fresh water,
as is the case for the divergent Trans Polar Drift. Upon export into
the convective gyres of the subarctic North Atlantic the low salinity
arctic outow acts in poorly understood ways to inuence convection and the global overturning cell.
Because of its importance in determining stratication within
the AO and its potential importance in governing deep convection
in the adjacent subarctic Atlantic, much effort has gone into
observing and modeling variability in freshwater budget of the
AO, both in terms of uxes into and out of the basin and storage.
The original estimate of Aagaard and Carmack (1989) was that
the EB held 12.2 103 km3 of liquid fresh water while the AB held
the larger volume of 45.8 103 km3, despite the fact that the sum
total of river inows to the EB is much larger than that to the
Amerasian. The total freshwater content at this time, including
shelf domains and sea ice, was found to be near 100 103 km3,
approximately equal to the freshwater stored globally in lakes.
The larger volume in the AB was attributed to the storage of PW
within the Beaufort Gyre. Subsequently, the retreat of the Pacic/Atlantic halocline front from the vicinity of the Lomonosov to
the AlphaMendeleyev Ridge in the late 1980s and early 1990s
(McLaughlin et al., 1996; Morison et al., 1998) would have necessarily reduced this volume. Morison et al. (2006) argued for a
return of the AO to pre-1900s hydrography, but did not place this
strictly in the context of freshwater storage. Proshutinsky et al.
(2002) advanced a numerical and a conceptual model to purport
that storage and subsequent release of freshwater from the Beaufort Gyre and export to the Nordic Seas could inhibit deep convention. Since then, a number of studies have discussed interannual
variability in AO freshwater content as forced by winds, increased
river inow and sea ice melt. Proshutinsky et al. (2009) examined
year to year variability in the southern Canada Basin and found an
unprecedented increase in freshwater storage of 25% during the
rst decade of the 21st Century. McPhee et al. (2009) and Giles
et al. (2012) also conrmed the increase in freshwater storage
within the basin associated with wind-driven convergence in the
Beaufort Gyre. Rabe et al. (2014) calculated a trend in freshwater
storage between 1992 and 2012 for the entire AO (for bottom
depths >500) of 600 300 km3 yr1. A decrease in salinity made
up about 2/3 of the freshwater trend and a deepening of the upper
layer the remaining 1/3. Time and space variability in storage and
pathways are discussed in papers by Newton et al. (2008), Morison
et al. (2012), and Korhonen et al. (2012).
In any discussions of the AO freshwater budget, its future state
and its role in biogeochemical processes, it is critically important to
identify the source of the fresh water, be it river, ice melt, direct
precipitation or Pacic inow, this is done through the judicious
use of geochemical tracers such as d18O, barium, N/P ratios and
alkalinity (Carmack et al., 2008; Yamamoto-Kawai et al., 2008,
2009). This is important biologically, because freshwater from
102
103
104
Fig. 9. Schematic sections illustrating the distributions of water masses and vertical biomass distributions along representative sections crossing the Arctic basins; inset
shows the location of sections: (a) from the Bering Sea across the Lomonosov Ridge and into the Greenland Sea; (b) from the Bering Sea and through the Canada Basin and
Canadian Arctic Archipelago into Bafn Bay, (c) from the Bering Sea across the Lomonosov Ridge and onto an interior shelf off Siberia. Colored circles represent average
biomass distribution across all basins of mesozooplankton (red) and macrobenthos (blue) with water depth (modied from Kosobokova, 2012 and Bluhm et al., 2011b). AW
Atlantic Water, DW Deep Water, HC Halocline, PW Pacic Water, SW Surface Water, SAT St. Anna Trough; white dotted line denotes the approximate upper boundary of the
homogeneous layer.
Fig. 10. Map showing overlay of expatriate zooplankton distributions with halocline circulation patterns and the Atlantic/Pacic halocline front (colors as in Fig. 5d). Only
records deeper than 500 m are shown. Data from Kosobokova (2012).
basins (Fig. 12a and b), but the depth of the maximum depends
on the seasonal state of the zooplankton community and the interplay between populations of migrating species. Regional variability
is related to the proximity of the Atlantic gateway area (Fig. 12a,
proles highlighted in blue), where advected populations of the
Atlantic copepod Calanus nmarchicus cause a biomass maximum
105
Fig. 11. Maps showing (a) zooplankton biomass (g DW m2) (modied from Kosobokova and Hirche, 2009; Kosobokova and Hopcroft, 2010) and (b) infauna (macrofauna)
biomass (g C m2) (modied from Bluhm et al., 2011b) for the two Arctic basins. Biomass is concentrated along the shelf breaks and in inow areas. Only records deeper than
500 m are shown.
Fig. 12. Vertical distribution of zooplankton biomass (mg m3) in (a) the Eurasian and (b) Amerasian basins. Blue are stations close to pronounced AW or PW inuence, red
indicates stations north of 85N, green shows the stations distributed elsewhere. Data from Kosobokova and Hirche (2009), Kosobokova and Hopcroft (2010), and Kosobokova
(2012).
106
penetrate into the AB due to their short, one year long life spans.
The AB receives hardly any allochthonous addition to locally
produced biomass, because the majority of plankton advected from
the North Pacic already dies off on the extended shelf of the
Chukchi Sea. Consequently, oceanic communities of the AB are
characterized by signicantly lower biomass compared to the EB
(Fig. 10).
Although there are pronounced seasonal variations of zooplankton abundance and biomass in the surface water layer of the Arctic
basins (Kosobokova, 1982, 2012; Ashjian et al., 2003), the vertical
zooplankton distribution pattern shown in Fig. 12 undergoes little
changes during the annual cycle. Year-round observation in the AB
revealed that abundance and biomass per unit volume at depths do
not increase substantially below 100 m even when key Arctic zooplankton species descend to overwintering depths, because they
are distributed over a wide depth range of several hundred meters
(Geinrikh et al., 1983; Kosobokova, 1982, 1983, 2012; Ashjian et al.,
2003; Darnis and Fortier, 2014). Density maxima remain within
the upper 0100 m layer throughout the year (Kosobokova, 1982,
2012; Ashjian et al., 2003).
Unique to the Arctic basins, the lower margin of the epipelagic
zone seems to be shallower than in the rest of World oceans where
it occupies the upper 200 m of the water column (Vinogradov,
1970). An abrupt drop in zooplankton biomass below 100 m
(Fig. 12) (Kosobokova, 2012), a noticeable upward shift of vertical
ranges of common mesopelagic species compared to elsewhere
(polar emergence, Kosobokova, 1989, 2012; Kosobokova and
Hopcroft, 2010), and a boundary between statistically distinct epipelagic and upper mesopelagic zooplankton communities at
100 m (Kosobokova et al., 2011) suggest that the epipelagic rises
up to depths of about 100 m in the Arctic. This phenomenon is
probably related to a combination of limited light penetration into
the water column due to low sun angle, presence of sea ice with
snow cover, strong stratication of the upper water column, and
overall low primary production concentrating fresh food in a thin
surface water layer (Zenkevitch, 1963; Harding, 1966;
Kosobokova, 1989, 2012).
The basin zooplankton fauna uses several mechanisms to cope
with the food limitation. In the surface layers, ice-associated and
upper water column pelagic crustaceans are closely tied to fresh
algal production as reected in their relatively low d15N ratios,
indicators of low trophic level (Iken et al., 2005). The deeper plankters use different food sources, because a considerable part of the
fresh organic matter is already converted into fecal material and
marine snow by ice-associated and surface-dwelling planktonic
herbivores and recycled the microbial loop within the epipelagic
zone before it sinks to depth (Olli et al., 2006). Consequently, carnivory and omnivory/detritivory are prominent feeding modes in
meso- and bathypelagic Arctic zooplankton communities
(Harding, 1974; Laakmann et al., 2009; Kosobokova et al., 2002,
2011). A number of these deep planktonic carnivores and detritivores have modied gut passages as an adaptation for more efcient digestion and assimilation of nutritionally poor and
digestively resistant food. Sigma-shaped (e.g. in the copepod
Aetideopsis rostrata) or substantially widened guts (e.g. in the copepods Scaphocalanus acrocephalus and S. polaris) permit longer
retention of food inside the gut (Kosobokova, unpublished). One
of the most striking examples is the copepod Spinocalanus antarcticus which possesses a strongly elongated and looped midgut,
enabling it to digest marine snow and organic coating of
tiny-sized mineral particles melting out of the ice and sinking
through the water column (Kosobokova et al., 2002). Vertical niche
partitioning, whereby closely-related zooplankton species occupy
different depth ranges to reduce resource competition, is another
adaptation used by the AO deep-water plankters in their
2007;
5.2. Benthos
As for zooplankton, abundance and biomass also decreases with
water depth at the Arctic (and global) deep-sea seaoor (Soltwedel,
2000; MacDonald et al., 2010a,b). This decrease tends to be greater
for benthic macrofauna (infauna) than for meiofauna (Wei et al.,
2010). This trend has been interpreted as an average decrease in
metazoan size with increasing water depth (Klages et al., 2004;
Rex and Etter, 2010) and as increased importance of smaller organisms with increasing water depth (Pfannkuche and Soltwedel,
1998; Rex et al., 2006). Generally, the range of macrofaunal densities and biomass (mostly below 4000 ind m2 and <1 g C m2,
respectively, summarized by Klages et al., 2004 and Bluhm et al.,
2011a; Fig. 11) fall within the lower end of values reported from
the North Atlantic (Levin and Gooday, 2003). Meiofaunal densities
in the central Arctic under ice-cover are either lower than in the
global deep sea (Schewe, 2001) or on the same order of magnitude
as in other oligotrophic deep areas (<100 to >3000 ind 10 cm2;
Soltwedel, 2000; Vanreusel et al., 2000; Hoste et al., 2007;
Grska et al., 2014). Despite the general trend of decreasing density
and biomass with depth, however, faunal densities and biomass
may vary substantially in areas of similar depths, depending on
vertical in situ and advective carbon ux to the seaoor that is
related to ice-edge effects and vicinity to inow areas in the AO
(Schewe and Soltwedel, 2003; Bluhm et al., 2005, see also Section 6.3). Klages et al. (2004) concluded cautiously, based on the
relatively sparse biomass and oxygen consumption rates available
for the Arctic deep-sea benthos, that consumption rates are generally both low, though regionally variable with lowest rates in the
basins, and in agreement with those from other deep-sea areas,
but that vertical organic matter supply may be higher than
expected from calculations of vertical carbon ux from sediment
traps.
The food web at the sea oor of both Arctic basins is still poorly
described. We do know, however, that it is characterized by a high
degree of reworking of organic material resulting in four to ve
trophic levels (excluding marine mammals; Iken et al., 2005;
Bergmann et al., 2009; van Oevelen et al., 2011). Benthic procaryotes and, in the larger size classes, deposit feeders play a major role
in carbon recycling as reected in the dominance of procaryotes in
a carbon ow model in the EB (van Oevelen et al., 2011) and in
enriched d15N signatures of macrofaunal biomass-dominant
deposit feeders and those of their predators and scavengers in
the AB (Iken et al., 2005). Typical for the deep sea, suspension feeders are less common in the abyss of the AB (Bluhm et al., 2005;
MacDonald et al., 2010a,b) because of extremely small currents
(Timmermans et al., 2010), although this feeding type is more
prevalent in the more rapidly moving deep waters in Fram Strait
(Bergmann et al., 2009, 2011). Interestingly (and somewhat contrary to the above concept that smaller organisms become more
important with depth), some larger members of the benthic community in both Arctic Basins are apparently capable of quickly and
efciently ingesting and utilizing fresh material when it does
become available, which has been long been documented from
other deep-sea areas (e.g. Billett et al., 2001). The rapid respond
to food pulses kilometers under the Arctic sea surface was recently
conrmed from observations of holothurian (sea cucumber) consumers of fresh ice algal matter at the seaoor (Boetius et al.,
2013) and low d15N ratios (indicating low trophic level) in some
additional benthic taxa (Iken et al., 2005, Iken and Bluhm unpublished). In distant locations in both Arctic basins, the same
surface-feeding mobile holothurians, Kolga hyalina, congregated
107
108
Fig. 13. Map showing the distribution of some (a) pelagic and (b) benthic taxa occurring in the Eurasian and Amerasian basins. Taxa are identied in the legend. The
distributions suggest that the Lomonosov Ridge does not serve as a distribution barrier. Note that for many other species, even fewer distribution records exist for the Arctic
basins preventing a general conclusion on distribution patterns. Data from Bluhm et al. (2011b) and Kosobokova et al. (2011).
109
continental slope of the Beaufort Sea that coincides with the comparatively warm Atlantic layer (Suydam et al., 2005; Citta et al.,
2013). The occurrence of belugas along the western Beaufort slope
is also correlated with a well-developed Alaska Coastal Current
producing strong frontal features (Stafford et al., 2013) and these
authors suggest this mechanism would provide enhanced foraging
opportunities.
Mesoscale eddy formation provides another mechanism for off
shelf transport of material properties and biota. Baroclinic eddies
have long been recognized to populate the halocline of basin interiors (Hunkins, 1974; Newton et al., 1974); an early census suggests that eddies comprise up to one quarter of the surface of the
southern Canada Basin (Manley and Hunkins, 1985). More
recently, Zhao et al. (2014) examined ice-tethered proler data
deployed between 2004 and 2013 to carry out a census of mesoscale eddies within the AO halocline in the basins. They documented 127 eddies, 95% of which were anticyclonic, the majority
of which had anomalously cold cores and were observed in the
Beaufort Gyre (AB eddies) and the Transpolar Drift (EB eddies).
Such eddies typically have horizontal length scales on the order
of 1020 km and horizontal (azimuthal) velocities of order 10
25 cm s1 (Timmermans et al., 2010) and their formation has been
attributed to instabilities related to current/topography interaction
(DAsaro, 1988), frontal zone processes (Timmermans et al., 2008)
and shelf break jets (Pickart et al., 2005, 2013; Spall et al., 2008).
Offshore transport of shelf-origin waters by eddies of Pacic origin
in the upper halocline are associated with elevated concentrations
of nutrients, organic carbon, and suspended particles (Mathis et al.,
2007; Watanabe et al., 2012; Pickart et al., 2013). These authors
attributed these features to derive from the boundary current
along the edge of the Chukchi Shelf. OBrien et al. (2011) examined
sediment trap data from the slope off the Canadian Beaufort Shelf
and argued that eddy phenomena also played a major role in transporting sediments from shelf to basin. For the Canada Basin, eddies
in three different depth domains were recognized: (1) shallow
upper halocline eddies centered at 80 m), (2) lower halocline
eddies (200 m) and (3) deep eddies (1200 m) (Carpenter and
Timmermans, 2012). Eddies observed in the deep waters at depths
between 200 and 2000 m of both the AB (Swift et al., 1997;
Carpenter and Timmermans, 2012) and EB (Schauer et al., 2002;
Aagaard et al., 2008) had water mass properties derived from adjacent shelves. Velocities within deep eddy cores were found to
range from 225 cm s1, and thus appear to have the potential to
transport material properties within and below the Atlantic layer
and re-suspend particles where Atlantic water abuts the continental slope (Carpenter and Timmermans, 2012).
Sea ice, through its drift patterns, also transports large amounts
of particles from rivers and the shelves into the central AO (Eicken
et al., 2005). This transport mechanism is in fact thought to have
critically shaped the sedimentation regime in the AO in the geological record (e.g. Nrgaard-Pedersen et al., 1998). Particle types are
comprised not only of sediments and terrigenous carbon that are in
part region-specic in terms of their composition and can, therefore, be used as tracers (Dethleff et al., 2000), but also of fresh algal
material that provides an allochthonous food source for basin
fauna (Boetius et al., 2013).
6.3.2. Basin to shelf
Wind forcing is the principal mechanism driving basin waters
onto the shelf, either via upwelling favorable winds (generally
easterlies) bringing water from depth onto the shelf, or via downwelling favorable winds (generally westerlies) driving surface
waters onto the shelf. In the arctic wind forcing may take place
in regions that are ice covered, partially ice covered or ice free
(see Wadams, 2000, for discussion). Clearly, however, as sea ice
continues to retreat, more and more of the shelf break domain is
110
111
Table 2
Observed and modeled changes in the Amerasian and Eurasian Basins in the past few decades.
Layer
Sea ice
Decreasing salinity, increased stratication, deepening nutricline (McLaughlin and Carmack, 2010; Jackson et al., 2010)
Observed and modeled high variability in freshwater distributions and pathways (Newton et al., 2008, Polyakov et al. 2008,
Morison et al., 2012)
Very low nutrient concentration in Beaufort Gyre suggests no/
little increase in primary production is possible (cf. Codispoti
et al., 2013)
Small phytoplankton thrive (Li et al., 2009)
Decrease for small zooplankton dependent on authochtonous
production, increase/no change in large zooplankton beneting from allochtonous production (Hunt et al., 2014)?
Aragonite-saturation decreasing (Yamamoto-Kawai et al.,
2009)
Enhanced shelf-break upwelling (Carmack and Chapman,
2003; Tremblay et al., 2011; Pickart et al., 2013)
Interannual shifts in location of the Atlantic-Pacic halocline
front (McLaughlin et al., 1996; Karcher et al., 2012)
Warm pulse penetration into the basin (Shimada et al., 2004)
and temperature increase between 0 and <1 C (McLaughlin
et al., 2009; Polyakov et al., 2013)
More intense anticyclonic Beaufort Gyre after 2004 constraining the cyclonic ACBC that transports AW (Proshutinsky et al.,
2009; Karcher et al., 2012)
Geothermal warming (0.004 C per decade; Carmack et al.,
2012), no time series to document biological change
Halo-cline
Atlantic water
Eurasian basin
Remaining MYI limited to north of Greenland (Barber et al.,
2015)
Ice algal production from shelves transported to basins, where
fast vertical ux supplied food to benthic fauna (Boetius et al.,
2013)
Variability in freshwater content (Timmermans et al., 2011)
Nutrient concentrations not limiting, perhaps increase in primary production possible (Codispoti et al., 2013)
No change in primary production between 1995 and 2007
(Wassmann et al., 2010)
Increased frequency of fall blooms (ACIA, 2005; Ardyna et al.,
2014)
(modeled) increase in secondary production across both
basins, slightly higher in Eurasian Basin (Slagstad et al.,
2011, their Fig. 9)
112
8. Outlook
Throughout much of the 20th century the AO and its central
deep basins in particular was viewed as a small, remote, slowly
changing and relatively unimportant part of the global system. A
perceived need to catch-up for lost time now prevails, requiring
that we pay more attention to the pan-Arctic perspective and to
the importance of advection and physical/biological connectivity
and their joint roles for the basins. This paper summarizes, from
an interdisciplinary point of view, our current perception of how
the Arctic Basins are set-up and operating, and how basin oceanography and biology have recently (e.g. the last few decades) changed
(Table 2). The AO is subject to large amplitude multi-decadal variability and long-term trends (Polyakov et al., 2013), thus challenging interpretations of observed changes to climate drivers. These
authors state, however, that the exceptional magnitude of recent
high-latitude changes, both oceanic and atmospheric, implies an
irreversible shift of the AO to a new climate state (see also
Jeffries et al., 2013; Wood et al., 2013). What does this new state
hold for biota? A review of the climate change footprints in arctic
ecosystems by Wassmann et al. (2011) gives evidence that all components of the high-latitude marine ecosystem are being impacted
by global change, though most existing reports considered large
mammals and birds only, and mainly in shelf areas.
What, then, do we know? Concrete ndings from decade-long
time-series span from declines in sea ice extent and thickness
(Kwok et al., 2009; Stroeve et al., 2012, Barber et al., 2015) to
increasing river discharges (McClelland et al., 2006), with consequences that include the appearance of aragonite-undersaturated
(low pH) waters in the Canada Basin this past decade
(Yamamoto-Kawai et al., 2008, 2011; Table 2). Other changes
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