Download outstanding the plants sharply distinguished: is always - UvA-DARE

The littoral environment of rocky
and
between the sea
as a
border
land and between the sea and
the
the fresh
C.
shores
den
water
Hartog
Summary
have
Many ecologists
disagree among
In this
paper the
sensu
in
lato
regarded
of the
explanation
an
the littoral
Secondly
water,
in
about the
has
problem
has been
approached
been
‘submergence’ of littoral organisms
action
(wash, splash,
acting
in the littoral
waters
are
restricted
the influence
third
active
supralittoral
with
action allows
estuarine
some
importance
well-defined
species
and
that
do
show
of the
almost
zone
does not result
is
no
of the
one
where
zone,
by
common
species.
factors
beyond
are
the
air.
three
own
the
physical
Strong
wave
regarded merely
be
The
abiotic
Both
pattern.
their
the
wave
low-salinity
they
These
zonation
to
for
major
in
is contact with
eulittoral.
this has
and the fresh
sea
explanation
movements and
tidal
characterized
supralittoral;
still
they
the zonation.
the littoral
‘submergence’
the littoral
units,
have
they
both
organisms
part
effectuate
ecological
in the
between the
of the intertidal
lower
in the
place
approach
reasonable
a
salinity
not
and
cause
the Arctic waters and the Baltic.
It appears that
which
This
zone
approach gives
factor for these
In fact
survive
to
the border
zonation
which
the first
In
in
organisms
as
waters.
(competition),
stress
are
the factors
and the land.
sea
upper part
The vital
species composition.
eulittoral
environment is in
border
it.
In
this environment
environment,
could indicate
for
and the
rhythm.
of
the
This
organisms
and
different ways.
two
fluctuations
salinity
of the
supralittoral
is
zones
as
an
phenomenon.
The littoral
to
low-salinity
in
cause
biological
own
some
environment.
Most
tidal
eulittoral
the
features and their
interference
daily
factor, light,
and
spray)
the
to
of the
factors, together
peculiar
and
zonation.
in
between
has been considered
lato
brackish
a
area
of the littoral
problem
the
of the
of many eulittoral
‘submergence’
as
with
delimitation
the border
as
zone sensu
other words
themselves
occupied
themselves
which
that marine salt
has
few
very
beyond
other brackish
comparison with
the influence
species exclusively
the reach
of the
habitats
extends far above
of the salt
regular
bound
tidal
it,
to
rhythm
extremely
the tidal
the
reverse
creates
rich in
influence.
is the
conditions
species
In
the
This
case.
favourable
speciation.
I.
One
of the
most
and animals in
Introductory
outstanding
features of the
number of often
a
remarks
very
shore is the
arrangement of the
demarcated
sharply
Three main
zones.
plants
zones
have been distinguished:
i.
The sublittoral is always submerged except for its
sometimes
be exposed
under
exceptional
within die sublittoral is generally ascribed
change
in the
The eulittoral
2.
light quality
or
intertidal
with
zone
to
increasing
is
the
the levels of
sometimes
mean
mean
pattern within
movements
375
subjected
It is
is
mean
is chosen
regarded
to
occurrence
may
of belts
the tidal oscillations
usually defined
low-water springs (MLWS) and
the intertidal belt
The
depth.
high-water springs (MHWS)
and the various
margin which
upper
decreasing light intensity and to a gradual
zone
where emersion and submersion alternate.
extreme
circumstances.
as
the
as
as
the
high-water
its
upper
product
ecological side-effects coupled
zone
mark
and
between
(MHW);
limit. The zonation
of the vertical
tidal
with the tidal factor.
BLUMEA
376
3.
The supralittoral is
the
sometimes reached
by the
to
The
be
general
with
respect
with hydrographic lines, such
spring and
neap
tion could
not
the
as
a
is
tides (Colman,
1933;
at
communities
above
m
a
feet above
far above the
sheltered
on
the existence of
40—50
mean
shores and recorded
The
exposure
nature
of the substratum
parts of the intertidal
that
the
on
a
the
various
other
also play
higher
occurs
at
the
a
on
number of
coincide
to
action, the
a
nature
of the
aspect
etc.
shores several
while
zone,
'raising'
occurrence
belts
Fucaceae
the
on
to
up
of various Fucaceae
canaliculata
tmder the influence of
coasts
of the British Isles.
in the
higher
observed
was
Vilvordian limestone than
level between these
at
(1954) and Conway
important role, especially
the shore
cor-
recorded
(1903)
of Pelvetia
Powell
&
algal
the
from the Faeroes reaching
marked
places along
an
correla-
a
determining
slope and the
exposed
On
such
cases
In the south-western part of the Netherlands it
zone.
granite it
some
can
Pelvetia-belt is situated
while
basalt,
of the
Fair Isle and
wave
occur,
sea-level. Burrows, Conway, Lodge
on
In
intertidal. Borgesen
umbilicalis
sea-level. He also observed
exposed
on
for
be satisfied with the
their part in
of the intertidal
limit
community of Porphyra
mean
factors
have been found
play
zone, e.g.
obviously
are
to
in many other
factors
important.
very
one
in the flood water, the
matter
upper
coasts
(1954) also recorded the 'raising'
increasing
zone.
levels of high- and low-water mark during
modifying
factor in particular is
levels
causal
movements.
zone
the time of the day when the lowest tides
exposure
occur
for
complex
too
Doty, 1946), but
be found. Numerous
responding belts
5
the major
as
result of the tidal
the position of the various belts in the intertidal
The
exposure
supralittoral respectively. There is, however,
average
substratum, the quantity of suspended
belts
is
the major factors governing the eulittoral zonation
to
opinion the tidal factor
my
of the shore,
margin
of
degrees
zonation within this
further
the borders of the various belts in the intertidal
cases
c.
a
been accepted
that the littoral zonation is
statement
for
responsible
have generally
spray
agreement
In
pattern.
Its lower
spray.
short time. The various
a
zonation patterns of the sublittoral and the
no
1968
2,
limit of the supralittoral is generally indistinct.
upper
Light and
for
waves
to
No.
XVI,
under the influence of the
zone
considered
are
spray
VOL.
on
(den Hartog, 1959).
two
Another feature which certainly complicates the correlation of the biotic communities
with tidal levels is the seasonal shift of various algal belts. Borgesen
the
supralittoral
In
the
the
move
downwards in
of these vegetations
can
be concluded that
by
the
Bangia-Urospora-,
show
summer.
but also
observations have been confirmed
it
the
associations
Enteromorpha-Porphyra
limits
that
Netherlands I observed
while they
(1905) reported that
extension of Porphyra umbilicalis is much greater in winter than in
These shifts
shown
are
by their lower limits
by
Nienhuis
(den
not
the borders of the plant and animal belts in the eulittoral
zone
upper
These
1959).
In
(1964).
the action and counteraction of the various
general
modifying
will rarely show
factors
any
correlation with the tidal levels. Further the eulittoral, here meaning the intertidal
will rarely coincide
by Kylin (1918),
the effect of the
the borders of
exactly with
a
who defmed the
biological
was
the other side
conspicuous
(Sjostedt,
into the
physically
definition of the various
with
species
on
1928; Levring,
quite right when he objected
biotic belts
belts
a
against
tidal levels
the
one
1937,
the
according
to
zone,
in order
to account
in order
to
to
find
for
correlate
an
easily
side and the supra- and sublittoral
1940).
Lewis
procedure
defined eulittoral
direct
difficulty already recognized
Nevertheless, workers continued trying
detectable border between the eulittoral
on
was
'physiological high-water line'
modifying factors.
some
unit. This
and
half-year,
only by the
Hartog,
and Lewis
(unpubl.)
summer.
minima-,
in the winter
shift
upward
an
Blidingia
zone.
In
of
(1961,
trying
place
1962), therefore,
to
fit
of this he
their dominant organisms.
the various
proposed
An
the
important
C.
objection
Hartog:
this scheme is that
to
organism,
DEN
as
composition of
west coast
coasts
usually
only
According
of the Atlantic (Borgesen,
because
which
found
intertidal
are
supralittoral
terminology).
sheltered
on
For
the
as
united
eulittoral
organisms
littoral belt defined in this
accompanied
organism'.
Along
It is
by
not
the Atlantic
tideless Baltic
Sea
the
to
enough just
it is
refer
to
a
p.
well-developed supralittoral
In
supralittoral
is
is
supralittoral
of
a
same on an
zone
same
of
an
gradients
sheltered
coast.
(Du Rietz,
tidal
a
exposed
exposed
as
This feature
individuality
In such
paper
the littoral
cases
I will
may
be the
to
two
and
In
key
I will refer
approach
This
separate
is
secondly,
as
as
it will be
a
I
think
units
enough
a
produce
to
Waern, 1952).
here the
number of species
indicates that in the
habitat with
more
or
part of the eulittoral
upper
explanation
of littoral zonation.
between
a
a
'littoral fringe'
'mid-littoral zone'
that he noticed himself that biolog-
Although
as
problem
Lewis did.
as
biological
1940;
coast
supralittoral) and
means
to
'sublittoral
negligible and
produce
in the
an
and
Laminaria
Levring,
sheltered
separate, it
them
to
the
to
term
limit of
the second place
as
action
can
the
upper
this definition needs
organism'
1964) distinguished
zones.
be kept
complex will be regarded
Gerlach, 1963)
a
on
action
produced by the tidal factor
ical differences exist between the
In this
and
wave
in his
I have various
organisms.
action is strong
wave
the
regarded
the eulittoral belt but in the
to
1925, 1932;
coast
coast
coast
(comprising the eulittoral communities).
together.
'littoral
wave
separate units,
proposed
from the
place
be regarded
to
the communities usually regarded
have sufficient
In the first
coast
Moreover, in all his diagrams Lewis (1961,
(containing
marine
of all
as
('midlittorar
reduced. The fact that the zonation pattern of
very
essentially the
less the
zone
sheltered localities along
very
coasts
number of species
a
Lewis
species of Fucus and
to
the tideless Baltic
000). Along
1959);
tidal
on
Borgesen (1905),
permanently submerged species.
a
quite
extending
zone
concepts
that the supralittoral and the eulittoral have
(see also
like
vesiculosus is confined
Fucus
coasts
the
way.
definition of the
a
be regarded
and supralittoral
limit
upper
cannot
contains
fact he,
as
the sublittoral
be
the also
den Hartog,
1912;
extension of the eulittoral
objections
a
Jonsson,
sublittoral species which
shores
In
coasts.
upward
an
'littoral' and he defined it 'biologically'
to
dominant
same
be mentioned. The floristic
respects from its composition
many
1905;
numerous
of exposed
zone
merely
zone
in
and eulittoral
supralittoral
the supralittoral
have the
may
may
low-water mark.
near
Lewis the
to
377
Fjord (Sundene, 1953) and along
(Kylin, 1918) differs
in the first mentioned areas it contains
shores
of rocky
dissimilar environments
this belt in the non-tidal Oslo
non-tidal Swedish
are
littoral environment
example of this the Fucus vesiculosus belt
an
along the tidal
The
in
may
my
be
the
opinion
useful
two
zones
consider them
to
the 'littoral complex'.
of the littoral zonation in
the transition between
regarded
as
sea
two
ways,
first
and land (see also
the transition between
sea
and fresh
water.
THE LITTORAL ZONE SENSU LATO AS
2.
If the intertidal belt is considered
land then the
the
most
periods of high
as
A BORDER
the intertidal
zone
is
the marine regime, while during low tides the
habitat because it is
then subjected
to
In the transition zone between these
of
a
population
penetrating
377
the
composed of
zone
from the
a
sea
is
obviously
same
zone can
conflicting regimes
mixture of
the
one
the tidal
sea
factor,
as
LAND
and the
during
completely submerged and subjected
the influences governing
two
on
SEA AND
representing the border between the
important ecological factor
water
ZONE BETWEEN
very
be
regarded
as
a
to
terrestrial
the adjacent land habitats.
one can
tolerant and
expect
very
side and from the land
the
occurrence
resistant
on
species,
the other side.
BLUMEA
378
when
Usually
not true,
of the
however,
XVI,
No.
sufficiently stable
is
zone
of a number of specialised
composed
This is
transition
a
VOL.
well
forms,
terrestrial
poorly
forms. This is
for
in
represented
for
true
a
of the intertidal
number of
and Lichina pygmaea,
are
Rhodophyceae,
the
Almost
all
the
intertidal
Orthoptera-Gryllidae,
penetrate
Schuster
restricted
in
occur
the
the sublittoral.
could
(1962)
of
the
origin, but
Araneae, Acarina,
to the intertidal
eulittoral
as
well
found
essential
are
that
not
find
any
correlation
In the
air
supply
are now
submersion in
supralittoral,
even
they
for
species
In fact all
exposed shores.
faithful
of
a
They
environments.
marina,
but bryophytes,
belong
invade
the
contains
even
a
upper part
tolerate
de
of the
Lichina
Virville
eulittoral
domain is still
so
It is
in
sea
1931; Du
be drawn here is that the intertidal
to
supralittoral
sea
and
land;
the primary
this transition
a
number
more
are
is also
or
on
less
the home
in all other terrestrial
Verrucaria
maura,
Caloplaca
fern Asplenium
A few of these
coasts.
water,
Further
species
e.g.
Lecanora.
Rietz,
1925,
marinum also
supralittoral species
true
the supralittoral
of which
some
of the lichen
The last
four
can
genera
genera
1930, 1932, 1938, 1940;
1932;
Hiibschmann,
von
no matter
zone cannot
zone
to
whether they
this
be regarded
zone.
as
The
are
con-
the transition
intertidal belt and the
consists of the
together.
factor causing
probably
even
by
&
more
A.
(1958)
that
the variation
the littoral zonation pattern has
action, regarded by Southward merely
A.
Schuster
constrictum, and Ramalina siliquosa,
restricted in their distribution
The conclusion reached by Southward
T.
supralittoral
important that the majority of the species,
clusion
between
levels
of the
that the marine influence in this terrestrial
or
are
tidal
structure
Several of these
spray.
supralittoral
confinis,
Ramalina, and
of marine
zone
The
sheltered
on
striking, however,
terrestrial origin,
with
the
still quite
are
supralittoral species (Davy de Virville,
Fischer-Piette,
&
Degelius, 1939).
on
water.
from
the small
supralittoral.
Xanthoria, Rhizocarpon,
true
Arthropods
of submersion.
period
completely absent
maritimum, and
of the
species
groups have
levels high in the supralittoral
to
complex.
are
e.g.
short-lasting submersion
a
have also developed
1957;
the
number of resistant and salt-tolerant terrestrial species,
Parmelia, Lecidea,
Davy
species
Collembola,
and
of these
organisms
better than in fresh
helicopis, Rhizocarpon
L.
Schistidium
e.g.
the faithful
to
mainly lichens,
are
actophila,
Lecanora
genera
the number of cavities in the substratum. These
but 'rise'
which
species
None
of these
for their existence
coasts
the communities of the littoral
to
number of terrestrial
zonation
of marine origin in the
species
to
tolerant to long periods of desiccation and obviously
are
sheltered
on
—
mucosa
strictly confined
to
obviously dependent
were
much
Verrucaria
last-mentioned
two
saline soils.
respiration during
sea water
obtain the salt and moisture
necessary
eulittoral
the
and
Arthropods.
are
above the highest flood level, there
of species of marine origin which
are
Only
one
marine
distinct zonation in their
pattern of occurrence
a
governed by
needed
not
is uncertain
Pseudoscorpionidea, Chilopoda,
non-marine
show
between the
that
found, however,
for the
tolerate
they
on
quite
Chlorophyceae,
e.g.
Diptera-Chironomidae belong
environment.
as
substratum. Their penetrationto lower levels is
cavities
and
Although these organisms
He
algal vegetations.
or
forms
Coleoptera-Staphylinidae,
completely
which
species
and
Phaeophyceae,
the eulittoral. Most of the littoral forms of terrestrial origin
are
there is
algae
The intertidal lichens,
of terrestrial
certainly
It is
well-developed but mainly derived from
they might be derived from terrestrials.
which
zone.
animal groups. The origin of the few intertidal Cyanophyceae
most
own,
dominant. The terrestrial element is
Among
respect.
every
element of its
an
the transitional environment.
to
rocky-shore population
The littoral element is
species.
it contains
adapted
obvious that here the marine element is absolutely
extremely
1968
2,
as
a
modifying factor,
important when looked
Stephenson (1949).
'In
a
to
upon
marine
from
area
with
the
no
in
water-level
is
be reconsidered. Wave
is
just
point
as
important.
of view
tide and
It is
expressed
no wave
action
C.
there would thus be
produce
we
third
a
the
zone,
littoral
corresponding
zones
each with its subzones. If
once,
The
Hartog:
den
to
related
littoral,
if tidal action is added
the
water,
the effects of
to
zonation is
that the belt of Verrucaria
investigators
within the tidal limits
occurs
sometimes
find that
can
Given
cause
the littoral zonation. As
ideal
an
communities will
animal
without
coast
move
action then
wave
soon
as
action increases these communities become
in the water-level.
Generally
the effect of the tidal
only the effect of the
generally
governed
by
cooperating
two
It is
known,
vertical
that there
however,
general lines similar
that of
to
water movements and
where the differences between
is the
pattern
fluctuations.
same
these
can
areas
the Fucus
be ascribed
1959).
Hartog,
surface
to
the
dwindles
occurs
as
Gulf, where
also the
of
far
it
will
and
wave
of the fluctuations
action modifies
wave
modify
movements
between
sea
movements
and land is
and the
serratus
Uschakov,
can
periods
as
more
move
and
remains
more
does
not
wave
are
of
down
to
usually
deeper
as
and
water
in
water
is
of the littoral zonation.
is
water
tides; during such wind-
may
become uncovered.
of low
salinity.
The
gradually
also be of importance
may
(den
low-salinity
deeper
and the number of littoral species
vesiculosus reaches the treshold of the Bothnian
In the northern Atlantic
1952).
the northern
on
sublittoral.
of Prince Edward
coast
below low-water
Similar
1959) and Novaya Zemlya
restricted
(1955)
low-salinity
that in
to
algae;
to
it
extremely important
Spray and fluctuations
as
it
Island
and that
'submergence'
(Guryanova, Sachs
surface water,
low-salinity
the
refers
coast
mark,
of
1954,
of
Murmansk
records
the
presence
not
to
along
phenomenon
Remane
feature, however,
level
to
in the
the Baltic the influence of
depth (Waern,
in the
been found
well.
the
zonation
littoral species in the sublittoral has been recorded. T. A.
exclusively
the
changes
cause
to
general
correlated
evident. Parallel with the decrease in salinity the
m
10
found that
explained by
not
factors
permanently submerged. This phenomenon
It has been observed for both littoral and sublittoral
This
but is
vesiculosus-belt
vesiculosus-belt extended
have
the
small,
very
to
and the Kattegat,
Fjord,
reach the Baltic, but Laminaria saccharina (L.) Lamour.
1930), Greenland (Lund,
groups
the Oslo
Europe
of surface
deeper
move
of melting ice. The
into
animal
of
part of the Fucus
Kiel Bay, and Fucus
occurred
species
be
tides
coast
be coupled
not seem to
Skagerrak,
in this sublittoral environment
the Fucus
Scotia)
a
serratus
occurs
occurrence
littoral
This
tidal
to
exposure
independent
zone
one
during spring
where the pattern of the zonation is
but does
and low
west
periodic influence
Stephenson (1954)
A.
(Nova
Fucus
Fucus
out.
the
In the southern Kattegat and
becomes
water
the
coast
In
spray.
high
on
some areas
are
tidal
a
serratus- belt
Fucaceae and Laminariaceae
&
of
degree
intervals strong off-shore winds
periods
decreasing light intensity
still
the
factors,
places
involved the plant
shores the tidal
considerably exceed the fluctuations due
can
induced low-water
In
that
as
irregular
At
which
level,
more
exposed
In such
submerged
sheltered shores
the transition
Thus
action.
wave
on
and
on
shores
exposed
on
('spray').
action
as
while
movements,
more
that
can state
one
and other
(1961)
the vertical tidal oscillations
When the
the shore.
up
of the
importance
Lewis
spray.
becomes
action
wave
The
by
are
waves.
all; but naturally
at
supralittoral
terrestrial lichens
salt-tolerant
tides.
which is
maura
of the
at
tide,
no
interface between air and
an
marked.'
more
sheltered shores without
on very
tide
zones
action but
amplitude
any
the observation made
supported by
wave
average
action and of
wave
and made
strengthened
action factor is also
wave
in
the
to
steady
therefore have all essentials of intertidal zonation without
We
379
supralittoral and infralittoral
our
add considerable and
we
shores
of rocky
environment
(Zinova,
in the
especially
waters many
littoral
has been observed
'brackish-water
&
1929).
species
in
many
submergence'.
species.
with respect
to
of the water-level
a
possible explanation
are
compound
factors
BLUMEA
380
having
in
a
a
salinity
common
stratified brackish
and
submergence
VOL.
component.
water
itself rules
such
z,
1968
oflittoral
degree of submergence
the Baltic shows
as
an
being
causal factor in the
a
zonation pattern of the littoral species which exhibit this. The fact that the
be involved in the littoral zonation
to
seems
littoral complex
transition
a
as
ZONE SENSU LATO
THE LITTORAL
3.
between the
zone
AS
FRESH
Dahl (1959) has pointed
the habitats
first step in becoming
the
emersion
inhabitants. The
ephemerous
taken
as
a
measure
of
development
importance for
I have taken
zone
an
by
exposed
the
to
the intertidal
cycle,
a
be
that 'at this stage
wrote
marine
or
not,
more
zone
particular
cannot any
even
be
This is often
factor of
we
than
can
step is the
next
zone.
by
period of
the various
to
to
supposed that the
in the life
pelagic phase
species should be called
a
is
species
taken
already
During
which they colonize the eulittoral
completely confined
of the
area
of conflict between the
if it lives
longer
on
an
sea
1964, 1967) by regarding
and fresh
water.
means
that in
increases.
can
to
is
intertidal
more
population
less
or
subjected
the
subjected
is
to
salinity. During
constant
zone
is
the marine
the
higher levels possibly
must
beable
As
to
a
must
be able
to
withstand
for several days. Further they
of desiccation
result
period
the direct influence of the weather.
to
withstand similar periods in which dry off-shore winds
the salinity
or
strong
concentration
withstand the sudden shock of becoming submerged
their metabolism
re-adjust
to
reason
at
desiccation.
Finally they
and
and
to
cause
a
weather the species in the intertidal belt
wet
hours,
equally be able
insolation
has
population
same
the
period
which generally
rain for several
the
I have classified it
the alternation of submersion and emersion.
the submersion
water
of emersion the
every
taxon
zone.
these influences in order
tolerance. Dahl
ecological
is in fact
species
this line of thought again (den Hartog,
up
as
about
During
surface
again
to
to
of
one
from marine organisms. The
of the main brackish habitats. The 'brackish' character of the intertidal
one
brought
must
extent
AND
coast.'
intertidal
This
of such
adapt
marine animals. Dahl
many
definitely whether
oceanic
as
special
a
by the loss
accompanied
say
of their
to
BETWEEN SEA
certainly have been
zone must
have evolved
of the population
part
upper
ZONE
that penetrates into the intertidal
influences of the weather; they have
the
and the fresh-water regime.
BORDER
brackish-water
poikilohaline
a
sublittoral species
every
A
sea
salinity factor
for considering
WATER
that the eulittoral
out
where brackish-water species
the possibility
opens
organisms
obvious correlation with salinity
of desiccation
possibility
the
out
The
No.
XVI,
regard the intertidal
the marine concentration. There is
to
zone
as
environment with unstable
an
thus
salinity
conditions.
The level of
mean
high-water springs, however,
influence. Therefore, in
was
treated
too
my
former
paper
simply. The supralittoral
on
this
zone
is
not
the
upper
is also
a
form of
out
wash, splash,
and
spray.
by the tidal
movements
The aerial transport of
of consideration as the influence of this factor
be taken into
of aerial salt
account
certainly
for which the
in
is
a
matter
sea
1).
and by
wave
action, in the
salt in the wind has been left
be felt
so
far inland that it
cannot
discussion of the littoral border environment. The availability
responsible
supralittoral
can
the
part of the border environment
between the conflicting marine and fresh-water regimes (Fig.
The marine influence is exercised
limit of the marine
subject (den Hartog, 1967)
is
too
for the distribution of several maritime
saline,
or
for the
unexpected
occurrence
organisms
of characteristic
C.
Fig.
1.
Diagram
den
Hartog:
The
littoral
of the conflict situation between
environment
the
sea
of rocky
and the fresh
shores
water
on
381
a
rocky
shore.
382
BLUMEA
littoral species
of
at
VOL.
distance from the
some
No.
XVI,
sea,
Rhodochorton
e.g.
The fresh-water influence is
as
rain,
dew,
snow,
exercised
the various forms of
mainly by
Further there is the surface
etc.
erably enriched by dissolved materials from the ground
this run-off
mineral compounds
also contains
water
bird droppings and rotting seaweeds. It
of the vegetation,
1932)
and
by birds
it favours the
as
small gullies
cut
biotope',
place,
from
inhabited by
turbellarians,
On
a
a
where
with
as
clearest
to
exposed
As
a
and
since
belts, according
am
modifying
use
well
for
rain coincide
stenohaline forms
they
can
short periods of
exposure
algae occurring
in this
of this belt
brackish
must
be
mean
a
very
destructive effect
of the
position
as
this
the
seems
low-water
on
when low
the
neap
small.
water
algal growth,
conditions.
low-water
neap
tidal cycle.
As
the salinity fluctuations
belt
mean
usually small. However,
are
and
mean
high-water
a consequence
are not
is
Most of the
extreme.
very
neap
of the relatively
rather euryhaline and penetrate into the polyhaline
are
very
high-water
mean
short periods and
neap
and
mean
high-water springs
of the time is fully exposed
most
salinity
and several of them
euryhaline
mean
high-water springs
less continually under the influence of splash,
part of this belt is
reached by the
lower supralittoral
storms.
belts,
are
are
very
widely
large.
to
The
the
algae
distributed in other
waters.
Above the level of
Finally
be
can
increasing fresh-
the actual
indicate the various
various weather conditions. The fluctuations in the
5.
environment
of estuaries.
submerged only for
average
to
all.
and the effect of weather conditions is
have
every
The belt between the levels of
greatly,
at
vegetation
border
finally determine
to
die under such
covered and uncovered during
4.
any
is
and
archiannellids,
discontinuously
the
This biotope
versa.
The vegetation is limited
low-water springs and
mean
short periods
very
The belt between the levels of
is
vice
isopods,
biotope,
'shocks' take
salinity
that the relation with the tidal levels is only
aware
factors which
the tidal levels
visited
follows the
eulittoral
special
sudden
two
if there is
to
This usually
a very
fauna of amphipods,
result the fluctuations in salinity
section
to
(1925,
Du Rietz
to
from
composition
frequented by birds.
not
semidiurnal tidal cycle the littoral
a
the
on
According
the rocks.
rise
species,
consid-
which it has flown. Besides
supralittoral rocks frequently
those
to
precipitation,
usually
demonstrate the discontinuity in the gradient of the fresh-water influence.
only
pouring
on
which is
marked effect
faithful indicator species.
The belt between the levels of
is
over
maximum and the other
a
are
number of
a
it is the
way
to
few Enteromorpha
a
nevertheless I will
belts,
3.
streets
organic substances originating
a
tidal cycle
every
run-off,
of Prasiolaceae.
giving
zone,
characteristic
of which
some
minima and
coast
during
influence. Although I
indirect,
have
trickle along
may
littoral
minimum
a
very
subdivided into
water
growth
composition
water
through the
the 'shock
one
in
quite different
are
Blidingia
can
Degelius (1939) the lichen vegetations
Further, phreatic fresh
2.
in the
purpureum
Flushing.
such
1.
iosB
2,
extremes
a
(1928). The salinity
during periods
is
due
to
now
excessive
completely
belt where the
This belt is
sea
a
belt
which is
more
or
and aerial salt transport. The lower
and then. This belt is the 'swell zone'
or
defined by Sjostedt (1928). The salinity in this belt fluctuates
being
here
salinity
there is
as
there follows
spray,
spray
exercises
the 'storm zone'
in this belt is
of bad weather.
on an
evaporation
dependent
or
excessive
or
climatic
its influence
upper
average
on
only seasonally
supralittoral
very
rain
low, but
fall.
The
circumstances.
in
may
the
sense
increase
and
of
during
Sjostcdt
considerably
G.
This
the
to
zone
The
littoral
beyond the reach
of
occurrence
of the
sea water
a
few times each
and then for
year
marine sublittoral and the
complete without reference
partly
low low-water
extreme
completely uncovered
or
emersion
is
period
small, but this does happen
and then.
Although
storm
and
fully
short periods. The chance that its vegetation
very
an
383
would not be
This strip is either
and fauna will be damaged during such
now
shores
of rocky
disturbance belt between the levels of
a
springs and ofmean low-water springs.
only
environment
of the littoral zonation between the
exposition
maritime
Hartog:
den
zone
would expect
one
and the maritime
the influence of the
conditions
exceptional
similar
a
as
zone,
will
spray
disturbance belt
the force of
with it, this is
vary
neutralized by the fact that
are
the border between
at
is also
storms
a
the
not
case
the maritime
the
variable factor
very
the effects of
as
zone
normally
is
under the influence of aerial salt.
When the littoral
fresh
water
it
Further, there
conflict
areas,
zone
are
indirectly
or
from
element is
In
water.
the estuary
Kiitz.
only represented by
a
surprising
sea
Further, there
are
Rhine,
and fresh
water,
species which
seem to
occur
of
this
is
group
submergence
depth.
In his
very
in
study
brackish
on
water,
sea
e.g.
fresh-
a
and fresh
where there is
in the section with
species
an
average
open
salinity
salinity
to
to
the eulittoral and the
descending
supra-
for the littoral border environment.
is considered it becomes obvious
with several of the species showing
in brackish
submergence
between
areas
is
character.
flows off,
water
total absence of
Scheldt,
there
the fresh-water element in the benthos
be confined
be faithful indicator
heterogeneous
for such
Blidingia
1967, 1968, unpubl.).
If the reaction of these various littoral species
that
and
and
involved originates
ephemerous
very
or
conflict
Meuse,
few animals which
seem to
a
representation
of the various
between 0.3 and 1.8 °/
Cl' (den Hartog,
00
littoral, and which
has
water
are
Generally
species.
the fresh
as
sea
its flora and fauna.
among
supralittoral, where phreatic
The weak
of the rivers
communication between the
not
in the
regular feature
a
between the
area
brackish-water species. Examples
are
precipitation, and thus
occur
Cladaphora glomerata (L.)
water
conflict
a
and various Enteromorpha
oxyspermum,
True fresh-water species
is
these
fresh-water element. This is in fact
directly
as
preponderant
number of species, mostly of marine origin, characteristic
a
and undoubtedly
minima, Monostroma
no
lato is considered
sensu
that the marine element is
appears
the sublittoral
into
Remane
water
to
a
the feature
considerable
(1955) distinguished three
different types of submergence.
a.
It
In the
can
case
of'upper-limit submergence' only the
be caused
species
to
greater
upper
by several factors. Decreased salinity
depths
where the
physiological requirements.
salinity conditions
In transition
the change from the regular tidal
such
areas,
movements to
limit of
are more
as
a
species is involved.
of the surface
water
drives the
in agreement with their
the Skagerrak and the
Kattegat
irregular wind-induced fluctuations of
the water-level excludes several sublittoral species from the lower levels of the eulittoral.
Unfavourable temperatures
species,
b.
'Basal submergence'
depths when
are
may
also
especially along the border
it
a
downward shift of the
area
limit of
a
a
species
descending
to
greater
the brackish-water environment. The species involved
usually eurybiontic species and
in brackish waters,
species. This
extension of their distribution.
upper
of distribution.
involves the lower limit of
penetrates into
tion from stenohaline
cause
of its
appears
Examples
to
are
therefore, they have less competi-
be the deciding factor for the downward
given
in table
1.
BLUMEA
384
VOL.
Table
LITTORAL
SPECIES WHICH
XVI,
No.
2,
1968
1
BASAL BRACKISH-WATER
EXHIBIT
SUBMERGENCE
Chlorophyceae:
Enteromorpha
intestinalis
(L.)
Rhizoclonium
Link
F.
Enteromorpha prolifera (O.
MUll.) J. Ag.
Harv.
riparium (Roth)
Ulothrix
flacca
Ulothrix
pseudoflacca
Ulothrix
subflaccida
Thur.
(Dillw.)
in
Lejol.
Wille
Wille
Rhodophyceae:
Hildenbrandia
purpureum
(Lightf.)
Rosenv.
'Complete submergence' involves the descent of both the
c.
of
Nardo
prototypus
Rhodochorton
a
species. This feature is shown by
These algae
usually
dwindle
where this
out
Examples
follow the layer
layer reaches
in table
given
are
many
of
a
water
SPECIES
with the
depth where light
same
salinity
becomes the
conditions and
limiting factor.
2.
Table
LITTORAL
and lower limits
upper
littoral species in the Kattegat and the Baltic.
WHICH
EXHIBIT
2
TOTAL BRACKISH-WATER
SUBMERGENCE
Chlorophyceae:
Monostroma
Tellamia
grevillei (Thur.)
contorta
Wittr.
Batt.
Phaeophyceae:
Ascophyllum
Elachista
nodosum
fucicola
Fucus
distichus
Fucus
serratus
Fucus
vesiculosus
Pilayella
Ralfsia
Aresch.
L.
L.
littoralis
verrucosa
Spongonema
(L.) Lejol.*
(Veil.)
L.
(L.) Kjellm.
(Aresch.) J. Ag.
tomentosa
(Huds.)
Kütz.
Rhodophyceae:
Callithamnion
Dumontia
Gigartina
Plumaria
stellata
too
F.
(O.
(Stackh.)
has
not
Ag.
Mtill.)
Lamour.
Batt.
elegans (Bonnem.)
'Submergence'
by
C.
scopulorum
incrassata
Schm.
been recorded for
algae
the fact that in the estuarine environment the
large
and the
salinity
stratification
earlier (den
Hartog, 1964, 1967)
Baltic
algae and animals
many
salinity
than
they
can
loaded with suspended
This is easily explained
unstable for
most
algae.
I
live permanently in
an
estuary.
Further,
material light penetration
is
as
water
the
with
waters
strongly
a
are
generally
have already shown
that under the rather stable salinity
can
withstand in
too
in estuaries.
daily salinity fluctuations
conditions of the
much lower
of estuaries
reduced,
are
average
usually
preventing
any
considerable descent of the algae. Finally, because of sediments the substratum in estuaries
C.
is
den
The
Hartog:
littoral
often unsuitable for epilithic algal growth.
some
It has
animals from sand- and sediment bottoms
environment. The
shows
Miill.
'basal
only
be
to
do show
pointed
1965), while the polychaete
(Vader,
385
out,
submergence
Lindstrom offers
amphipod Bathyporeia pilosa
of'complete submergence'
shores
of rocky
environment
an
excellent
certain
of
range
reduced capacity
Submergence
water.
fluctuations and
salinity
for competing
in brackish
The latter are
is
water
in the wave-dashed
exhibited by
not
zone,
or
supralittoral
in the
spray
terrestrial domains do
exhibit
It is in fact remarkable
inhabit the higher
belts
rhythm, but rather
to
in brackish
water
is
on
that almost
the
they
can
not
are
are
given
restricted
in Table
to
the littoral
CONFINED
TO
LITTORAL
are
exposed
and do
gave
to
not
the
the
groenlandicum J.
Rosenvingiella
Suhr
in
constricta
COMPLEX
Ag.
(Setch.
&
(Rosenv.)
Gardn.)
Silva
Silva
hartzei Rosenv.
Urospora
Urospora penicilliformis
Aresch.
(Roth)
Eulittoral:
minima
Blidingia
(Nag.
Capsosiphon fulvescens
Monostroma
ex
(G.
oxyspermum
Kütz.) Kylin
Ag.)
Setch.
&
Gardn.
(Kiitz.) Doty
Phaeophyceae:
Supralittoral:
Waerniella
lucifuga (Kuck.)
Kylin
Eulittoral:
*
*
*
*
L.
Fucus
spiralis
Fucus
virsoides
Mesospora
*
Nemoderma
*
Pelvetia
*
Petalonia
Species
which
(Don)
macrocarpa
(L.)
zosterifolia
to
be
Ag.
(Feldin.)
tingitanum
canaliculata
need
C.
den
Hartog
nov.
comb.*)
Schousboe
Dene
(Reinkc)
submerged
&
Thur.
Kuntze
now
and
then in undiluted
sea
water
daily tidal
spray.
Jessen
Rosenvingiella polyrhiza
examples
exhibit submergence
Blidingia marginata (J. Ag.) Dangeard
stipitata
as
typically 'littoral'
to
Supralittoral:
Prasiola
a
add the lichens and the
Chlorophyceae:
Monostroma
by
water
(1955) had already
3
THE
nor
of the
3.
Table
SPECIES
zone
species,
zone
He
water.
all the species which
where
shore,
a
from the fresh-water and
the irregularities in the tidal rhythm and
A list ofspecies which
in brackish
submergence
zone
the rotifers, and the Halacaridae. To these I
ohgochaetes,
musci.
not
Remane
zone.
observed that the forms which have invaded the littoral
the
all essential
stable environment.
a
These all remain in the
species.
not at
partly specialists adapted
partly extremely eurybiontic species with
with stenobiontic forms in
considerable number of eulittoral
fluctuations,
F.
submergence'.
the species which submerge in brackish
to a
example
Nereis diversicolor O.
It is apparent that the tidal alternation of emersion and submersion is
to
however, that
in the estuarine
to
survive.
BLUMEA
386
Eulittoral,
but
also
Bifurcaria
*
Cystoseira myriophylloides
*
Elachista
*
scutulata
2,
1968
Sauv.
Aresch.
(Sm.)
velutinum
Himanthalia
No.
Ross
bifurcata
Herponema
XVI,
pools:
in
*
*
VOL.
(Grev.)
S.
elongata (L.)
J.
F.
Ag.
Gray
Rhodophyceae:
Supralittoral:
Bangia fuscopurpurea (Dillw.)
*
Porphyra
Lyngb.
linearis Grev.
Eulittoral:
Bostrychia scorpioides
*
*
arbuscula
Callithamnion
tetricum
Catenella
*
Caulacanthus
*
Ceramium
*
Ceramium
*
*
*
*
*
ustulatus
Nemalion
(Mert.)
welwitschii
Chauv.
lanosa
Ag.) J.
(Veil,
(L.)
Porphyra
leucosticta
Porphyra
purpurea
*
Porphyra
umbilicalis
Rissoella
verruculosa
*
Duby
Silva
Batt.
Ag.
Foslie
in
With.)
Batt.
Tandy
Porphyra elongata (Aresch.)
*
*
in
(Kiitz.)
(Esper)
tortuosum
helminthoides
Gray
Kiitz.
(Rupr.)
spinulosuin (C.
Polysiphonia
F.
Batt.
shuttleworthianum
Lithophyllum
S.
(Dillw.)
deslongchampsii
Erythrotrichia
Gelidium
(Dillw.) Lyngb.
(Lightf.)
repens
Mont.
(Huds.)
Callithamnion
Kylin
Thur.
(Roth)
(L.)
in
Lejol.
C.
Ag.
J. Ag.
(Bertoloni) J.
Ag.
Cyanophyceae:
Supralittoral:
Calothrix
scopulorum [(Web.
Entophysalis
Lyngbya
deusta
confervoides
[C.
Mohr)
&
(Menegh.)
Dr.
Plectonema
battersii
Schizothrix
calcicola
Symploca
atlantica
Born.
Gom.
Lyngbya semiplena [(C. Ag.) J. Ag.]
tenerrimus
Ag.]
Gom.
Ag.]
Lyngbya majuscula [(Dillw.) Harv.]
Microcoleus
C.
D.
&
Gom.
Gom.
Gom.
[C.
Gom.
Ag.]
Gom.
Eulittoral:
*
*
Rivularia
mesenterica
Rivularia
bullata
(Thur.)
& Flah.
Born.
[(Poir.) Berk.]
Born.
&
Flah.
Lichens:
Supralittoral:
Anaptychia
fusca
Bacidia umbrina
Caloplaca
(Huds.)
aractina
Vain.
Bausch
(Ach.)
(Fr.)
var.
marina
Hayrén
Caloplaca granulosa (Miill. Arg.) Jatta
Caloplaca
marina
(Weddell)
Caloplaca scopularis (Nyl.)
Caloplaca
Catillaria
Lecanora
thallincola
(Weddell)
chalybeia (Borr.)
actophila
Lecanora
Zahlbr.
Lettau
DR.
Mass.
Weddell
hehcopis (Wahlenb.)
Ach.
Sandst.
Lecanora
leprosescens
Lecanora
poliophaea (Wahlenb.)
Lecanora
riinicola
Lecanora
salina
H.
Lichina
confinis
Physcia
subobscura
Ramalina
H.
Ach.
Magn.
Magn.
(Miill.)
C.
Ag.
Nyl.
sihquosa (Huds.)
A.
L.
Sm.
Degelius
&
Flah.
C.
constrictum
Rhizocarpon
Rinodina
salina
Roccella
Verrucaria
shores
of rocky
environment
387
Malme
DC.
(L.)
Wahlenb.
ceuthocarpa
Verrucaria
littoral
Degelius
fuciforniis
Verrucaria
The
Hartog:
den
Wahlenb.
maura
symbalana Nyl.
Eulittoral:
Arthopyrenia
*
Lichina
orustensis
Erichs.
(Lightf.)
pygmaea
C.
Ag.
Verrucaria
erichsenii Zschacke
Verrucaria
microspora Nyl.
Verrucaria
mucosa
Wahlenb.
Ascomycetes:
Eulittoral:
*
Didymella
balani
Feldm.
(Winter)
Bryophyta:
Supralittoral:
Schistidium
maritimum
B.S.G.
(Turn.)
Ferns:
Supralittoral:
Asplenium
L.
marinuin
Angiospermae:
Suprahttoral:
Crithmum
Inula
maritimum L.
L.
crithmoides
Lebel
Spergularia rupicola
x
) Mesospora
Rev.
364—365;
In his
Algol.
on
paper
'Les
species Ralfsia
described
overgrown
indeed
that the
reported
marines
macrocarpa
(Aresch.) J. Ag.
Hartog,
de la Cote
in fact
but
also
'.
All
Although
material
and the
for
to
these
of the thallus
Mesospora
of Bot.
clatural
belt,
e.g.
where
as
it is
of
the
growth
means
elements
of
paraphyses,
features
of these
(see Feldmann,
possible
to
'species'
of this
was
crustaceous
to
1937, p.
to
brown
of the
epithet
'absolument
as
Soc.
(1935)
algae
does
by
The
preparation of
the
as
a
name
from
macrocarpa
of R.
that
not
only
at
Ralfsia
separation
a
between
(Article
the
of the
Ralfsia
indispensable
does
macrocarpa
to
the base
du
mediterranea.
cross-sections is
Ralfsia
he had
verrucosa,
the presence of small
permit
type
verrucosa
macrocarpa
Mesospora
Ralfsia
satisfactory
his earlier
caracteristique
of
not
that
Ralfsia
not
have
70 of the Intern.
Mesospora
is the
nomen-
of this.
species only
a
few
they
become
seem
fully exposed
ecological specialists. They
occurrence
are
to
be confined
to
to
the lower levels of the intertidal
the air and littoral pools it is improbable that they
probably restricted
specific
in the eulittoral of
to
the eulittoral
eulittoral;
some
such
Rhodophyceae,
can
as
as
a
result of
com-
perhaps be given
for
Callithamnion arbuscula,
and Ceramium shuttleworthianum. Elachista scutulata and Herponema velutinum
epiphytes
eulittoral. Cystoseira
reason
26
Himanthalia elongata and Bifurcaria bifurcata. As these species inhabit both places
C. tetricum,
are
to
the line of demarcation
similar explanation
petition with the sublittoral dominants. A
the
and
paraphyses,
be characteristic
observe
267).
of R.
similar
very
of
sample
a
pedicellate sporangia,
regarded
appeared
was
and the
of these elements
one
The transfer
Bull.
Feldm., ibid.
recorded
(1937)
mixtum, viz.
original description
Phaeophyceae. Therefore,
select
Feldm.,
macrocarpa
mediterranea
Feldmann
a
on
In his
sporangia
of cross-sections
crustaceous
Nomencl.).
consequence
Of these
of
possible by
rejected
Code
part
Alb£res'
The insertion of the
sporangia.
differentiating
the mode
it is
the identification
be
are
the base of the
the middle
on
macrocarpa
at
des
mediterranea.
but that it differed in the sizes of the unilocular
pedicellar cells
Ralfsia
—
Mesospora
—
had been based
by Mesospora
structure
comb.
nov.
10,pro parte.
f. 40—41.
263—267,
(1937)
9
algues
den
211—213, PL
(1931)
22
Lejol.
(Feldm.)
macrocarpa
Hist. Nat. Afr. Nord
ex
in this
of Himanthalia
myriophylloides
case
for restricting
competition
is
elongata,
only
and
known
are
with the sublittoral
its distribution
to
such
an
for
that
reason
from pools in the
Cystoseira species
unusual
biotope.
limited
upper
is
to
the
part of the
probably
the
VOL.
BLUMEA
388
No.
XVI,
Among the high-littoral and supralittoral species
of species which
the
water or
spray,
contains
group
depend
group
and the other of species
occur
The other
nitrogen-fixing Cyanophyceae.
Fucus
fluctuations in the salinity but for
Among the animals of this
to
the
of the
and which
group
seem to
some
can
they keep their
them
opening
larvae
period
occur
of 'stenohaline' eulittoral
The
of
group
as
period
a
adapted
are
with the
regard
during
of this
to
stenohaline marine species
into account, then it
to
escape
water
are
the
littoral
true
the effects of the instability of the habitat
them as indicator
regard
seems more correct to
group
salinity they could be regarded
brackish-water species. However, when the various mechanisms developed by
as
the
metabolism.
in brackish
submerge
not
surroundings,
of the thalli of Nemalion
structure
water
the
e.g.
feed. Their pelagic
they
unfavourable effects
escape
to
tolerance with
great
a
to
gelatinous
survive.
the periods of
protective function also. All representatives
species
which do
species
they show
contact
every
of submersion when
molluscs. The
among
and Rivularia probably has
species, and
closed, avoiding
marine. Similar mechanisms
strictly
are
emersion
valves
the short
during
enormous
water to
be stenohaline,
to
this
Pelvetia canaliculata,
as
barnacles Balanus balanoides (L.) and Chthamalus stellatus (Poli). During
emersion
in
the lichens and the
e.g.
need undilutedsea
be considered
even
penetrate
which can tolerate
helminthoides,
The first
coast.
Many species
species such
contains
one
of the flood
salinity
marine
fully
coast
every
be recognized,
can
groups
their metabolism,
some reason
group
two
lowering
and Blidingia minima.
and Nemalion
spiralis, Porphyra umbilicalis,
a
confined
with the air for
contact
to
along
Bangia fuscopurpurea
e.g.
on
which
species
some
freshwater,
into
be quite insensitive
seem to
1968
2,
species
are
some
taken
of the littoral
border environment.
The fact that the great
of the
majority
littoral and the uppermost part of
action
As
on
the
both
zonation.
contact
one may
supralittoral this
species do
with the air is just
certainly
of preponderant
significance.
must
the
For
As
light
exercise also
are
restricted
are
accentuates
is
and fluctuation of the
wonder whether
to
the supra-
the importance of
show
water
submergence
as
level have
is the
salinity alone
factor has been accepted
not
such,
cause
wave
and
but just the bare fact
in brackish
the factor aerial
salinity
a
of littoral
water
indicates that
important for them. In the eulittoral the salinity
as
importance
contact
seems
to
factor is
be of less
the causal factor for the zonation in the sublittoral this factor
influence in the eulittoral during
some
The influence of the light
on
action
wave
common,
supralittoral
littoral species
of speciation.
the factors
component in
that the
process
true
the eulittoral
on
the clarity of the flood
the intertidal zonation depends
water.
the
on
period of submersion.
the tidal difference and
Thus salinity, air ('the terrestrial influence'), and light
the three major abiotic factors
which
dominate the littoral
border
environment
(Fig- 2)-
4.
BIOLOGICAL
STRESS
AS
AN
ACTIVE
FACTOR
IN THE
LITTORAL
BORDER ENVIRONMENT
The
joint
action of the
explain the often
ment.
very
sharp
three abiotic factors salinity,
Without the action of
a
fourth
species would have much wider
would find
a
zonation. It is
not
not
factor, biological
ranges
series of intensely mixed
This factor is
air, and light is insufficient
to
demarcations between the belts in the littoral border environ-
on
populations
always fully appreciated
stress
or
competition, the
various
the shore and instead of clear-cut belts
which
and is
gradually
merge
usually omitted
mentioned in the work of T. A.
&
A.
one
into each other.
in studies
Stephenson (1949),
on
nor
littoral
in the
C.
Fig.
2.
Diagram showing
den
Hartog:
the influence
The
littoral
of the three
environment
abiotic
of rocky
major
shores
factors in
the littoral
389
complex.
BLUMEA
390
works
(1957), Southward (1958),
of Doty
in his
(1964),
Lewis
biotic factors
VOL.
the zonation pattern and rather
on
100
play
On
extremely important
an
the
At first diatoms
of
by patches
lactuca L.,
first
belts,
be
an
is the
any
obtained
a
with
years,
of
Fucus
example
&
however, the normal
well
biological
stress
however, quite
5.
THE
Although
I
between them
It does
not
water.
Only
in brackish
perhaps
zone
a
a
is
view.
soon
followed
Ulva
these species
becomes differentiated into
lower belt where
a
germlings
The various
of Fucaceae
species
at
grow
Entero-
and small
random
found Pelvetia canaliculata low
even
on
Ascophyllum nodosum. (Plate 1).
on
the
was
reinstated.
may
vegetation
Conway (1946),
have
to
rare
sometimes prevent
OF
can
any
THE
species
it contain
occurrence
There
are
single exception.
no
species
is
EULITTORAL
together
the differences
them
separate units.
as
action, particularly of
wave
is correlated with fluctuations of the
which
It is,
proportions.
greatly underestimated.
AND
maintaining
which show
species
In the
the zonation
on
1959).
and eulittoral
warrant
of wind and
product
whose
Lodge (1948),
work is needed before the role
SUPRALITTORAL
to
influence
be evaluated in its real
the suprahttoral
a
of the intertidal
pattern
Jones (1948),
significance for belt-forming
sufficiently essential
is
experimental
few of the species occurring in the
water.
time
a
this
(L.) Grev.,
Although
purpurea.
soon
zonation pattern
too
in littoral zonation
have discussed
does
competition
(1956), and Koumans-Goedbloed (1965).
Southward
INDIVIDUALITY
contain
level. Neither
I
snails
by patelloid
obvious that its
are
one
there it is
in the eulittoral.
are
compressa
of the barnacle Balanus balanoides
of the fact that much
typical supralittoral
The
P.
They
stones.
taken by littorinid snails (den Hartog,
is
aware
as
Chlorophyceae.
Lodge (1950),
place
pattern its
the
F. serratus, and
vesiculosus,
that
agree
support
usually takes place
on
vegetation.
for
Netherlands where Patella vulgata L.
of
After
well-known from the studies by
Burrows
data which
fine
succession
brownish slime
numerous.
of zonation,
The influence of browsing
am
must
zonation pattern often becomes disturbed
some
Ulothrix flacca, Enteromorpha
this pioneer
in
appear
trace
few
a
most
early spring settlement
is
I
it is apparent that
belt dominated by Blidingia minima and
upper
the establishment
belt
I
distributed the vegetation
randomly
shore, together
Within
forming
appear,
compressa
without
In
Netherlands, where the
maintenance,
Blidingia minima,
to
Rhodophyceae
I
algal belts.
influence of
possibility of
the
to
belt-forming.
Pilayella littoralis, Porphyra umbilicalis, and
seem
morpha
the
role in
newly laid dike slope the following
a
two
of the
coast
of dike
result
a
at
some
% effective. However, from recolonization experiments
Along
mentions the
shores',
reluctantly refers
notice the action of competition in well-balanced communities,
not
must
as
1968
2,
(1961), and Davy de Virville (1964).
Lewis
book 'The ecology of rocky
competition delimitating the lower borders of
does
No.
XVI,
spray.
water-
in brackish
'complete submergence'
supralittoral show 'basal submergence'
confined
are
to
undiluted
A number of salt-tolerant terrestrial
sea
species
water,
with
penetrate
the
from its
but marine invaders from the sublittoral are absent. Many of the
upper side,
species occurring
in the
The eulittoral is
wash,
one
the
the
form of
degree. Several species
with corresponding
mergence'
are
zone
peculiar
to
this
zone.
of the fluctuations
of the components of the wave-action factor.
marine element in
to some
supralittoral
essentially
sublittoral species
are
confined
to
which
of the water-level and the
It contains
are
able
the eulittoral because
to
a
well-developed
tolerate emersion
they
cannot compete
forms in the sublittoral. A number of species show
in brackish
water.
Further
a
considerable number of species,
'complete subnot
exhibiting
C.
Hartog:
DEN
brackish-water submergence,
littoral environment
The
strictly confined
are
Although several species of terrestrial origin
generally
this
to
The
part of the eulittoral
extremely small.
They
which
all
are
and Rhodochorton
the supralittoral
insufficient
for
reasons
has
are
water.
are
in the eulittoral are restricted
6.
in
supralittoral
e.g.
and
the eulittoral
is
Hildenbrandiaproto-
permits eulittoral species
action
wave
the form of reduced
supralittoral species
the
regarding
to
WITH
COMPARISON
be ascribed
merely
as
to
specimens
—
an
to move
and that
under such conditions,
occur
supralittoral
THE
mixed
seem
impoverished
to
fringe
the interference by the effects of
ESTUARINE
When the littoral border environment is
richness in species peculiar
border environment,
includes
the
Rhizoclonium riparium,
and fluctuations of the water-level.
action
only
one
species which
In fact
sea
transitional character.
a
to
exposure
of the eulittoral. These features have
wave
with undiluted
coasts
salt-tolerant terrestrial species
occurring
common to
ubiquists,
often
—
of eulittoral and
populations
me
391
purpureum.
The facts that increasing
into
shores
zone.
upper
The number of species
typus,
to
occur,
absent. A considerable number of species
of rocky
the
it is
to
number of
only
occur
in estuaries
not to
an
estuary, which is also
exclusive
species
are
confined
the estuary but
to
with other brackish habitats its
compared
In
Fucus ceranoidos L.
algal species,
they belong
striking.
ENVIRONMENT
characteristic
a
surprisingly
it is
to
its intertidal
to
zone
or
its
small,
and
Most of the
(den Hartog, unpubl.).
supralittoral.
the estuarine extension of the littoral border
environment.
In the discussion of the littoral
action
have exercised
must
In
movements.
in its widest
zone
sense
much greater influence
a
it became obvious
on
the speciation
the estuarine environment tidal action is the
that
wave
than the tidal
important factor,
most
and its influence extends much further than that of the salt. In the littoral border environment
just the reverse is
true.
the reach of the regular
this
respect
seems
areas
This could indicate that the influence of marine salt
tidal rhythm
it
is
to
be devoid of
It
ON
NOTE
to
it
(Caspers,
1955; den
Hartog,
border environment is
a
coinciding
the
The
with the
sense
term
eulittoral
coast.
joint
zone.
distinguished
reason
the
proposed,
The 'hygrohaline
The terminology
term
confusion. It
area
used by
littoral,
may
as
term
expresses
exactly
applied the
'hydrohaline'
can
fact better
in
and
the
in his work
on
'hydrohaline zone',
an
zone' of Du Rietz
and their 'aerohaline zone' is
the part of the terrestrial
the use of the
now
1967).
the various
to
(1925, 1932)
Du Rietz
eulittoral and sublittoral. Degelius
of eulittoral. For this
'amphihaline',
supralittoral,
cause
Du Rietz
In
brackish environment,
the zonation along the Baltic coast, and by Degelius (1939) in his work
west
of
TERMINOLOGY
terminology has already been applied by
a
out
favourable for speciation.
wonder whether this should be expressed in the names given
of the Swedish
in
A
exclusive
has been shown that the littoral
one may
belts. Such
on
species
conditions
border environment of fresh-water tidal
that the littoral
interesting
7.
and
creates
term
the lichens
more
as
less
better be avoided.
the character of the
Degelius coincides
same
or
hydrohaline
the maritime
with the
zone,
i.e.
that is under the influence of wind-transported marine salt.
Du Rietz
and Degelius
may
this has been used in several
also be useful
to
please those who object
senses
and thus
those who reject the use of the
may
term
to
sometimes
supralittoral
BLUMEA
392
and
prefer
other
the
to
dubious
a
presented here
terms
of lakes and other inland
complicate
too
were
to
waters
meat
2,
1968
new terms
has been
terms
charged
are not
with
any
'littoral terminology' I consider
current
fully equivalent alternatives,
of
set
unify the terminology for the
to
attempt
myself prefer the
I
be
to
Another
coasts.
No.
XVI,
like 'littoral fringe'. The
term
interpretation. Although
marine
VOL.
long
as
proposed by
zonation of the
Du
they
as
are
Rietz in
applied
in
1940
an
shore and that of the shores
sea
('limnological-thalassological
zonation system'), but these
approval.
any
REFERENCES
1905. The
F.
BGRGESEN,
Bot. Faeroes
E.
M.,
E.
Fair
Isle.
J.
E.
M.,
BURROWS,
on
BURROWS,
H.
GASPERS,
42:
coasts
& H. T. POWELL.
LODGE,
1950. Note
Stat.
with remarks
1954. The
the
on
phytogeography.
of intertidal
raising
algal
zones
of
zonation
62:
(1949)
Ver.
Limnol.
on
30—34.
613 —619.
12:
and animals.
J.
plants
and Fucus
of Patella, Balanus
inter-relationships
Port Erin
Verh. int.
des Elbeastuars.
of the intertidal
nature
the
on
Biol.
Mar.
Rep.
coast.
Limnologie
1955.
the Faeroese
of
283 —288.
& S. M. LODGE.
1933. The
COLMAN, J.
S. M.
CONWAY,
Ecol.
semi-exposed
a
algae vegetation
683 —.834.
2:
Biol. Ass.
mar.
U.K. 18:
435—476.
E.
CONWAY,
with
Isles,
Paris
DAHL,
1946.
sect.
17:
and
reference
special
raising
158: 752.
of intertidal
algal
exposed rocky
zones on
the zonation of Fucaceae.
to
—
Comm. 8e
Rapp.
of the British
coasts
Internat. Bot.
Congr.
133—134.
1959. Intertidal
E.
of Patella. Nature
Browsing
The modification
I954-
in
ecology
the
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