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Full text : Looking beyond stratification: a model-based analysis of the biological drivers of oxygen deficiency in the North Sea

2512

F.  Große  et  al.:  Looking  beyond  stratification

Biogeosciences,  13,  2511-2535,  2016

www.biogeosciences.net/13/2511/2016/

1  Introduction
Low  oxygen  (O2)  conditions  (concentrations  <  6  mg  O2  L _  1  ;
OSPAR-Commission,  2003),  often  referred  to  as  O2  deficiency, ­
  occur  regularly  in  the  North  Sea.  A  major  process  regulating ­
  the  seasonal  dynamics  of  bottom  O2  is  the  occurrence
and  duration  of  thermal  stratification  (e.g.,  Greenwood  et  ah,
2010;  O’Boyle  and  Nolan,  2010),  which  limits  the  vertical
exchange  of  O2  between  the  oxygenated  surface  layer  and  the
deeper  layers.  In  combination  with  events  of  enhanced  primary ­
  production,  and  the  subsequent  degradation  of  organic
matter,  this  favours  the  evolution  of  O2  deficiency  (e.g.,  Diaz
and  Rosenberg,  2008).  Although  the  northern  North  Sea  reveals ­
  strongest  stratification,  lowest  O2  concentrations  occur
in  the  central  North  Sea,  which  is  shallower  and  where  the
duration  of  stratification  is  shorter  and  shows  highest  year-toyear
  variability.  Thus,  one  can  argue  that  stratification  is  an
important  prerequisite  for  O2  deficiency,  but  its  severity  and
duration  is  controlled  by  the  complex  interaction  between  the
hydrodynamical  condition  and  the  biogeochemical  processes
involved.
The  North  Sea  is  a  temperate,  semi-enclosed  shelf  sea  adjacent ­
  to  the  northeastern  Atlantic  ocean.  It  has  an  average
depth  of  about  90  m  (Ducrotoy  et  ah,  2000)  with  northward
increasing  bottom  depth.  The  North  Sea  circulation  is  characterised ­
  by  a  cyclonic  pattern  mainly  driven  by  the  southward ­
  Atlantic  inflow  across  the  shelf  edge  defining  its  northern ­
  boundary.  Lenhart  and  Pohlmann  (1997)  showed  that
about  85  %  of  the  incoming  Atlantic  water  is  recirculated
north  of  the  Dogger  Bank,  a  shallow  area  with  water  depth
less  than  40  m  and  300  km  of  zonal  extent  at  about  55°  N,
2°  E  (Kroncke  and  Knust,  1995).  The  circulation  south  of  the
Dogger  Bank  is  governed  by  the  inflow  through  the  English
Channel  and  follows  the  continental  coast.  At  the  southern
tip  of  Norway  it  joins  the  Norwegian  coastal  current  leaving
the  North  Sea  at  its  northern  boundary.
Stratification  in  the  North  Sea  reveals  some  substantial  regional ­
  differences.  While  the  shallower  southern  parts  are
permanently  well-mixed  due  to  the  strong  influence  of  the
M2  tidal  component  (Otto  et  ah,  1990),  the  deeper  parts  north
of  54°  N  reveal  seasonal,  mostly  thermal  stratification  (e.g.,
Burt  et  ah,  2014;  Pingree  et  ah,  1978;  van  Leeuwen  et  ah,
2015).  Seasonal  haline  stratification  occurs  to  a  lesser  extent ­
  along  the  Norwegian  coast.  The  transition  between  these
permanently  mixed  and  seasonally  stratified  regions  occurs
gradually  (Pingree  et  ah,  1978).  In  consequence,  even  areas
relatively  near  to  the  coast,  which  are  affected  by  high  riverine ­
  nutrient  run-off,  often  reveal  stratified  conditions  at  subseasonal
  timescales  (e.g.,  Burt  et  ah,  2014).
In  the  1980s,  events  of  O2  deficiency  reaching  values  below ­
  3mg02L _1  occurred  regularly  in  the  stratified  southeastern ­
  central  North  Sea  and  in  the  German  Bight  (Brockmann
  and  Eberlein,  1986;  Brockmann  et  ah,  1990;  Rachor
and  Albrecht,  1983).  During  that  time,  the  problem  of  low  O2
conditions  in  the  North  Sea  reached  public  awareness  in  relation ­

  to  eutrophication  as  demersal  animals  died  across  a  large
area  due  to  these  low  O2  concentrations  (von  Westernhagen
et  ah,  1986).  Eutrophication,  or  in  other  words,  high  anthropogenic ­
  nutrient  loads  mainly  supplied  by  rivers  (Brockmann
et  ah,  1988;  Jickells,  1998;  Rabalais  et  ah,  2010),  may  raise
the  ambient  nutrient  concentrations  followed  by  an  increase
in  biomass  production.  Under  given  physical  conditions,  eutrophication ­
  thus  causes  an  enhanced  supply  of  organic  matter ­
  sinking  into  the  subsurface  layer  and  reinforces  O2  consumption ­
  near  the  sea  floor  due  to  bacterial  remineralisation.
Even  though  the  second  International  Conference  on  the
Protection  of  the  North  Sea  (INSC-2)  prescribed  a  50  %  reduction ­
  of  river  nutrient  loads  (inorganic  nitrogen  and  phosphorus) ­
  in  order  to  mitigate  the  effects  of  eutrophication
(de  Jong,  2006),  Fig.  1  shows  that  O2  deficiency  remains
a  persistent  problem  in  the  North  Sea  up  to  the  present  day.
According  to  Kemp  et  al.  (2009)  these  events  can  be  classified ­
  as  “persistent  seasonal”.  Low  bottom  O2  concentrations
may  cause  death  of  benthic  organisms  or  fish  eggs  as  well
as  avoidance  of  the  affected  areas  by  benthic  species.  Therefore, ­
  low  O2  concentrations  constitute  a  major  indicator  of
eutrophication  (category  3  indicator,  i.e.,  “evidence  of  undesirable ­
  disturbance”;  OSPAR-Commission,  2003)  and  concentrations ­
  lower  than  6mg02L _1  result  in  the  classification ­
  as  “problem  area”  in  terms  of  eutrophication  within  the
OSPAR  assessment  (Claussen  et  ah,  2009).  In  the  present
study  the  term  “oxygen  deficiency”  is  used  in  this  OSPAR
context  rather  than  “hypoxia”.  While  O2  deficiency  is  clearly
defined  within  OSPAR  by  the  6mg02L _1  threshold,  hypoxia ­
  refers  to  the  negative  impact  of  low  O2  concentrations
on  organisms.  An  overview  of  the  impact  of  hypoxia  on  marine ­
  biodiversity  can  be  found  in  Vaquer-Sunyer  and  Duarte
(2008).  Further  descriptions  on  the  ecological  disturbance  of
different  levels  of  low  O2  concentrations  are  summarised  by
Friedrich  et  al.  (2014)  and  Topcu  et  al.  (2009).
Despite  the  relevance  of  the  bottom  O2  concentrations  for
the  assessment  of  the  ecological  status  of  an  ecosystem,  O2
measurements  are  sparse  and  either  temporally  or  spatially
limited.  In  addition,  it  is  difficult  to  place  the  measurement  at
the  right  time  and  location  to  obtain  a  comprehensive  picture
of  the  duration  and  spatial  extent  of  summer  O2  deficiency
(Friedrich  et  ah,  2014).  One  way  to  address  this  problem  is  to
analyse  the  representativeness  of  available  data  with  respect
to  eutrophication  assessment  (Brockmann  and  Topcu,  2014).
Only  in  recent  years  continuous  measurements  for  the
North  Sea  have  become  available  by,  e.g.,  the  SmartBuoy
programme  of  Cefas  (Centre  for  Environment,  Fisheries  and
Aquaculture  Science,  UK;  Greenwood  et  ah,  2010)  or  the
MARNET  programme  (MARine  Monitoring  NETwork  in
the  North  Sea  and  Baltic  Sea)  of  the  BSH  (Federal  Maritime ­
  and  Hydrographic  Agency,  Germany).  These  monitoring ­
  programmes  provide  daily  time  series  of  O2  and  related
parameters  (e.g.,  temperature,  salinity,  chlorophyll)  in  different ­
  depths  and  allow  for  the  analysis  of  the  temporal  evolu-
            
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