the shetland isles: long-term observations on the subtidal marine flora

THE SHETLAND ISLES: LONG-TERM OBSERVATIONS ON THE SUBTIDAL
MARINE FLORA
IAN TITTLEY & W L L M F. FARNHAM
T m w , I. & W.F. FARNHAM.
2001. The Shetland Isles: Long-term observations
on the subtidal marine flora. Arquipdago. Life and Marine Sciences. Supplement
2 (Part B): 1-17. Ponta Delgada. ISSN 0873-4740.
The marine alga! flora of SuIlom Voe, Shetland, has been surveyed on three occasions in
twenty years (1973. 1983. 1993). To date 281 spec~eshave been recorded in Shetland. of
which 175 species occur in SulIom Voe, The subtidaI vegeration of the outer part of the voe
is charactensed by forest formations of the perennial kelp Laminaria hypcrborea on
bedrock and a species-rich eplflora on stipes and holdfasts. This community was stable with
time despite the wave-exposed conditions there. Tne subtidal vegetation of the inner,
sheltered, voe with a sea-bed of sediment, stone and sheEI, was characterised by the canopyforming annual Laminaria sacchantla / longicruris. Comparison of survey results revealed
this canopy and associated subflora communil!es to vary in abundance and extent w ~ t h
time.
fan Titsly { e m a i l : I. Tittlqv@nkm+ac.
uk), Depar~mentof Burany, The Natural Hisloty
Museum. London SW7 58D. VK. - Wtllia~rtF. Farnham, Inslitute of Marine Sciences,
Uuiversity of Portsmouth, Pormouth PO49 9LY, OK.
Although accounts of the algal flora of the
Shetlands go back almost two centuries (t.g.
Shetland is an archipelago of about 100 isIands in EDMONSTON
1809). studies have been sporadic
(1903~~
b),
the Atlantic Ocean lying between 60" and 61°N, and key works are those of B~RGESEN
B0RGESEN & J ~ N S S O N (1905). DEON (19631,
170 krn north of mainland Scotland and 300 km
southeast of the Faroe Islands. AIthough the IRYINE (1962, 1974, 1980) and IRVINE et al.
archipelago lies within the cold-temperate region, (1975). The marine algal flora of Shetland is
the sea does not freeze due to the warming typical of the species poorer cold temperate North
influence of the Gulf Stream (average surface Atlantic Ocean with 281 species recorded (50
temperatures are 4°C in winter and 12°C in Chlorophyta, 101 Phaeophyta, 130 Rhdophyta).
Sullom Voe had been scantly studied until the
summer). The Shetlands contain a mainland
which i s 85 km long and incised by deeply construction of Europe's largest oil terminal in the
& MCINmRE 1995)- Long-term
penetrating, fjordic arms (sea-lochs or voes); one 1970s (DUNNET
of these, Sutlorn Voe p i g . 11, is the subject of this surveiIlance of the intertidal biota of Sullom Voe
has been undertaken regularly since 1981
paper.
Sullam Voe is a 15 km long, narrow inIet of (MOORE et aI. 1995). Studies on the subtidal
vegetation were initiated in 1973 ( T ~ E etY al.
the sea where there is a gradient of wave-exposure
from extremely waveexposed to sheltered. A few 1977) as part of a wider study d the algae of
rivulets drain inta the voe but cause no significant Shetland (IRYINE 1974), and repeated in 1983
Y al. 1985) and 1993 ( T m E Y &
reduction in salinity. Sullom Voe has a range of ( T ~ E et
I997 - unpublished report).
subtidal habitat types, including rocky outcrops, FARNHAM
The 1993 algal survey of the Sullom Voe
extensive sediment areas of shell, stone, sand and
silt. and biogenic surfaces such as beds o f provided an opportunity to revisit previous study
sites, assess the current status of the subtidal
Modiolus (horse-mussel).
INTRODUCTION
vegetation, and make comparisons with previous
observations; some results are presented in this
paper.
dimensional graphs in which each point represents
a vegetation sample or quadrat. The distances
between the points in the graph are taken as an
indication of their simiIar~tyor difference.
METHODS
The surveys were carried out in July / Augusr
1973. July 1983 and August 1993. In the original
1973 survey, the principal features of the algal
vegetation of Sullam Voe were recorded by
sampling from quadrats along transects; these
transect studies were complemented by sampling
from 'spot-dive' study sites at pre-selected depths
( T ~ L EetY a[. 1977). As the original survey was
essentially semi-quantitative and descriptive,
transect sampling was not replicated; subsequent
reassessment was underraken in thc same manner.
Transects were relocated as accurately as previous
field data allowed. At each site a rope line was
laid from an accessible reference point at high
water mark to the subtidal region. The transects
were 200 m long and the rope forming the
transect line was weighted and marked at 10 m
intervals. Depths at these points were recorded at
noted dmes and adjusted to Chart Datum. At each
marked 10 rn interval, vegetation was cleared
from a 1 m x 1 m quadrat beside the transect line
and brought back to the laboratory for
identification. Where possible, whoIe stones and
cobbles were coIIected for examination of
encrusting algae. Laminarians were weighed wet
to assess relative abundance; ages of Lcrteinaria
Ityperborea plants were ascertained by sectioning
stipes and counting annual growth rings tcf. KAIN
1963). The results fnr four of the transect surveys
(Fig. 1) are presented below (for other sites see
T ~ L E&YFARNHAM
1997).
Detrended
Correspondence
Analysis
(DECORANA. a numerical, ordination method)
was used to facilitate comparison of quadrat data
from the three surveys. A database of species
presence in quadrats was prepared for the purpose
(in T m L E Y & FARNHAM1997). Ordination
arranges the vegetation samples in relation to each
other according to their similarity of species
composition and associated environmental or
temporal controls (KENT & COKER1992). The
results of an ordination are in this case two-
Fig. I . Sullom Voe showing itransects sites.
RESULTS
Inner Voe - South Ness
The transect lraversed an intertidal area of gently
sloping shingle, boulder and outcropping rocks
that continued subtidally and, at greater depths
(I0 m). gave way to a sea-bed of soft mud and
she11 debris.
Vegetation description and comparison
Seventy-nine species were recorded in the three
surveys (32 species in 1993,55 in 1983 and 36 in
1973); only a small component (11 species) of the
total flora was found on all occasions.
A1 shallow sublittoral levels the transect
crossed a canopy of mixed of Fucw serrarus,
Lruninaria digitotn, L hyperborea and 1,
sacchari~ldlor~gicruris (L. saccharinn and L
lottgicruris may be conspecific cf. SOUTH&
T~LEY
1986). From 30 m offshore (2-3 m
depth) the canopy was solely of L. saccharina /
lottgicruris (Fig. 2). In 1993 rhe Lmninaria
canopy extended to only 90 m offshore (9 m
depth), contrasting with 1973 when it was present
in all quadrats to 140 rn offshore (14 rn depth),
and 1983 when it was patchily present 80-130 m
offshore.
The subflora (underllora and epiflora)
recodcd in shallow waters comprised a diverse
species-assemblagebut differed in composition in
each survey. At 3C80 m offshore (2-8 rn depth)
only a depauperate underflora was recorded in
1993 compared with spenes-richer assemblages
of 1973 and 1983 (cf.TIITLEY
et al. 1977, 1985).
The principal aIgae in 1993 were, epiphytes on
Laminaria blades (e.g. Ecrocarpus siiiculosus,
Myrionema coru~lnae),and crustose forms such
as the Ag1ao:onia stage of Ciderin mvlt~ftrln,
Lthothamnioti glaciale and Pseudolithodcrrtla
extensunr on stones and shells. At 140-150 m
offshore only crustose species were recorded in
5993 in contrast to the foliose and filamentous
species found previously; in deepest waters at 2528 m (160-200 m offshore) vegetation was absent
in 1993 contrasting with prev~ousoccasions (cf.
T ~ E etY al. 1977, 1985).
The red alga
Phyllophvrn cri.~pnthat in 1973 and 1983 was
commonly present as extensive, detached, mats
120-200 m offshore, was not recorded in 1993.
Quadrat comparison
Ordination (Fig. 3) produced a single cluster of
points. The Iack of a clear linear pattern
successive points (quadrats along the transect)
reflects the patchy mosaic of vegetation crossed
by the transect. The first axis of the ordination
suggests a depth gradient; quadrats at depths of
tess than 10 m (characterised by Laminaria
saclzarirla / lungicruris) are positioned to the left
of 200, quadrats from deeper waters (often
containing Phyllophora crispn) are positioned to
the right. The second axis suggests temporal
differences with most 1973 and 1983 quadrats in
the lower part of the graph (below 200),and 1993
quadran in the upper part. The ordination does
not show the quadrats that in 1993 lacked algae.
200m
I
Abundance
-
= 2~~trn'
1
Fig. 2. Tmnsect 1, South Ness. Kite diagram showing
Larni~~ariusaccharina / longicruris extent and
abundance along the ttansect m the three survey years.
Inner Voe - Scarva Taing
The transect crossed an intertidal area of shingle
with rocky outcrops that gave way at shallow
sublittora1 levels (3 m depth) to a sea bed of
gravel, coarse sand and cobble. At 4-5 m depth
the sea-bed was of mud and sand which continued
to 100 m offshore. Beyond this point the sea-bed
changed to mud-shell debris and Modiolus beds,
and soft mud and shell debris at the greatest
depths investigated (25-30 m).
:
10
Xa
300
23
-
Z
m
.W
(II
-
200
*
I
I
rY
.-
r
920
I
1
II)
2
4
100
-32
1
'16
\
*rs,
,, .- - -,*45
I
L
#
%
1
-- --
a,?.
17
I
-_
1
'
$,'.
t
I I
"
1
,'
,a> 9
'
1
I
#
,#;',<A
om,
#$
;6
an
'', ,*
,'
,'
-34
'27
I
'b~;@
I----
-
\
--
''
-1I
*
+941
-'
,'
I
I
/'
I
.4+
=a!
-29
/
I
922
421
926
II
ah
-25
.Zt
E igenvalue 0.5506
Axis 1
1OD
200
300
t oa
Fig. 3. Transect 1. ordination of quadrat data. Numbered points represent quadrats (success~velyjoined for 1973
and 1993 transects). 1973, hatched line 35 (inshore) - 49 (offshore); 1983, points unjoined 16 (inshore) - 34
(offshore); 1993, continuous line 1 (inshore)- 15 (offshore).
Vegetation description and cornparisor1
Eighty species were recorded during the surveys
(42 in 1973; 64 in 1983; 18 in 1993) but only
eight on all wcasions.
In 1993 the principal kelp at sublittoral fringe
level was L saccharins / long icmris, contrasting
with mixed Laminaria spp, recorded in 1973 and
1983.
At greater depths a mixture of L
sacchurina / longicruris and L Izyperborea was
present on all occasions; L saccharinn /
Imgicruris was consistently the dominant species
in deepest waters (Fig. 4). Differences in extent of
L. saccharinn / longicruris canopy were observed
in each survey occasion (1973 to 110 m offshore;
1983 to 50 m offshore; 1993 to 40 rn offshore).
A diverse subflora of mainly red algae (e.g.
Phycodvs
rubens,
P!ty/lophorcr crispa,
Polysipl~oraiaeb~orlgatn,Pterosipl~or~
ia parasilica)
and the crustose brown algae Aglaozonia and
Pseudolitlzoderr~~a
extensum, was recorded in the
three surveys. Species composition differed on
each occasion and fewer species were recorded in
1993 than previously. The subflora extended to
only 80 m offshore in 1993, compared with 150 m
in 1973 and 200 m in 1983.
Om
Vegetarian description and comparison
Seventy-seven species have been recorded in the
surveys (47 in 1983,48 in 1983,46 in 1973) but
only sixteen species were recorded on all three
occasions.
In 1993 land 1973) Caminaria ddgitata and L.
Ityperborea successively formed narrow bands on
boulders at shallow sublittoral levels, contrasting
with 1983 when both were absent. L. saccharina /
longicruris was predominant in deeper waters
offshore on all occasions (Fig. 6). At the greatest
depths investigated in the present survey (20 m),
macrophytcs were absent. Laminaria succharina /
longicr~lrisbeds extended to 80 m offshore in
1993 compared with 190 m in 1983.
m
m
.-UI
Abundance
= 2~~lrn'
200m
Fig. 4. Transect 2, Scarva Tarng. Kite d~agram
showing Laminnria saccharina /longicntris extent and
abundance dong the transect in the three survey years.
Ordination (Fig. 5) produced a graph with a single
cluster of points. Quadrats from below 20 m depth
are positioned in the top left of the plot and those
from shallow water to the right side of the plot.
Quadrats from deep water and lacking in algae in
1993 are not shown.
-
Central Yoe Oil Terminal
The transect crossed a man-made boulder
foreshore that gave way in shallow waters to a
sandy sea-hd. At 4 m depth the sea-bed was of
mud and shell debris and changed to soft mud at
approximately 10 m depth. In deep water (15-20
m depth) the sea-floor was of soft mud and silt.
Abundance
1983
1993
Fig. 6. Transect 3, OiI Terminal. Kite diagram
showing I ~ m i n a r i a~accharina/ longicruris extent and
abundance along the transect in the h e survey years.
A subflora of Ulva lactuca, C h r d a $lam,
Ectocarpus spp., Sphacebria cirrosa and
Phycodqs rubcns occurred beneath the
h i n a r i a canopy on all occasions. The filiform
brown algae D~c~osiphotl foeniculaceus,
beyond 90 m offshore in 1993 while in 1983
Coccotylus mmcarus, Polysiphonia elongata,
Phyllophora c r i s p , and R. jloridula were
present, and in 1973 Polyides rotundus,
Des~narestiaanrlmta, Sphacelarin plunwsa and
Clwetomorpha rnelagonium were also recorded
(TI'ITLEY
et al. 1977, 1985).
Stic~os~pfron soriferus,
Spennatochnus
and Sauvageaugloia griflthsianus
recorded in 1973 and 1983 (cf. T ~ L E etY al.
1977, 1985) were not found in 1993. Flocculent
mats of Rhodothamniella floriduln occurred in
smaller amounts at 70 rn offshore in 1993
compared with 1983. Algae were not found
paradoxus
r
'On
X
-
,720
c
,
-3
*
5
#'
I
I
I
I
#
I
5
.-
)
t
I
I
#'
,'
I
,2,---------
1
I
I
I
-+--
pO'
/
1
3013 -
a
#
,
------- 5.26
*I
I
,#,!'
., I' :
E 1
,
''
I
I
: Ir
I
; .-I01
1
''.i
I
\
'21
128
'39
.I1
'35
Axis 1
*30
93
Eig~nvalue 0.57
'36
loo
200
300
400
500
Fig. 5 . Transecl2, ordination or quadrat data. Numbered points represent quadrats (successively joined for 1983
and 1993 transects). 1973, points unjoined 29 (inshore) - 41 (offshore); 1983. hatched line 9 (inshore) - 27
(offshore): 1993,continuous line l (inshore) - 8 (offshore).
Quadrat comparison
Ordination of quadrats (Fig. 7) indicated a depth
gradient with those horn deeper waters positioned
at higher values on axis 1; the arrangement of
successive transect quadrats in the graph is not
linear but meanders (from shallow to deep water)
reflecting the mosaic of communities traversed.
The second axis suggests a time series, with 1973
quadrats positioned at low values, 1983 quadrats
at middle values, and most 1993 quadrats at high
values.
N
t
0
0
3
m
300
-
$,
m
.w
N
u
200
-
3
16
826
113
'I0
*>I
"t",
.M
loo -
Axis 7
100
200
300
Eigenvafue 0.54
4 OD
Fig. 7. Transect 3, ordination of quadrat data Numbered points represent quadrats (successively joined for 1973
and 1993 transects). 1973. hatched line 26 (inshore) - 38 (offshore): 1983, polnts unjoincd I! (~nshore)- 27
(offshore):1993, continuous line L (mshore) - 10 (offshore).
Outer Voe - Grunn Taing
The transect crossed a wave-exposed intertidal
sea-shore of shallowly shelving bedrock which
extended to 5-41m below low water level ( I 00 rn
offshore). Eeyond this point were sporadic areas
of boulder, stones and gavel over bedrock, and
furthest offshore, parches of sand and small
boulders over bedrock.
Vegetation descriptiotr and conrparison
Fifty-nine specics have been recorded In the three
surveys (41 In 1973, 32 in 1983, and 39 in 1993).
Twenty species have been found on all occasions;
seven are unique to this site.
Beyond a narrow inshore band of L, digirata,
the principal kelp specics on all occasions was
La~rrirlnrlrrIryperlrorea which formed an extensive
stand of large plants to 200 rn offshore (Fig, 8). In
I993 plants were aged at (5)-6-17) years old.
Patchy growths of L snccharirtn / longicr~rris
grew among the L. l~yperhorencanopy along the
entire transect in 1993, compared with only
beyond 100 m in 1973 and 1883. Another kelp,
Succorltiw polyscfiides, was found occasionally
in the three surveys, where sand is temporarily
washed away from the bedrock.
The subflora in all surveys comprised (Table
1) a species-rich assemblage on Laminnria
hyperborea stipes, and a species-poorer
assemblage on bedrock and stones beneath the L
hyperborea canopy.
which a dear depth- or time-series could not be
discerned.
DISCUSSION
Fig. 8. Transect 4, Oil Terminal. Kite diagram
showing Laminaria hyperborean extent and abundance
along the transect in the thee survey years.
Table 1
Subflota a! Grunn Taing
Epiflora on Latnirraria hyperborea
Audouinelh purpurco
Cladophora mpestris
Cryptopleum mmsu
Derbesia marina
Dermdolitkon pusrurularum
Desmorestk viridis
HaraldiophylIwn
bommaisonii
hmentaria clawllosa
Mem branoptera alata
phycodrys-rubens
PIocomin~t
cartilagineum
Polysiphonia stricla
Ptilota gunneri
P~ilothamnionpluma
Rhodolnela confervoides
RRodophyIlis divadcuta
Sphcelaria mdicans
U a d e r f l m on M m k ody
Hildenbrandia ru bra
Peyssonnelia dubyi
Litbothamnion glaciak
Pkymalolithon
nrgulosum
Subflora on be&&
and 1, hyperborea
M~mbmnoplemdata:
Ptilata gunneri
Phycodlys rubem
Ulua lactuco
Po!ysiphania stricta
Quadrat comparison
Ordination (see T
~ & FAUNHAM
Y
1997,p.87)
presented a single tight cluster of quadrats in
There exists an extensive b d y of knowledge on
the sasonality, long-term occurrence, change in
abundance and depth distribution, of Laminaria
spp. around the coast of Scotland (e-g. WALKER&
RICHARDSON
1955; WALKER1957). Laminaria
occurrence around Shetland was less known until
recently tcf. CONNORet al. 1997; T
~ et al.Y
1977). The results of this Field survey agree with
the conclusion o f WALKER(1958) that the greater
abundance of Laminnria spp. occurs at depths of
less than 9 rn. The present and previous algal
surveys have revealed differences in laminarian
communities in the inner and outer parts of
SuIlom Voe. The extensive areas of subtidal
bedrock in the outer voe provide habitat suited to
the strongiy anchored, long-lived perennial
Laminaria hyperborea. The three surveys have
shown the species to thrive there despite the
wave-washed conditions which at an extreme can
cause considerable damage by snapping the stiff
stipcs (BIRKE~T
et al. 19981. WALKER (1958)
noted the dominance of L hyperborea on
extensive bedrock in the Orkney Islands. The
inner reaches of Sullom Voe have only a
restricted area of subtidal bedrock and therefore
little L hyperborea. However, the extensive seabed of sediment, stones and shells there, create a
suitable habitat for the fast-growing, opportunist,
annual L. saccharina 1longicruris and for filiform
annual brown algae that do not require firm
anchorage. Similar assemblages occurred in
sheltered fjords in the Faroes where L longicruris
grew on small stones and shells or even
unattached ( T ~ etYa]. 1982). With its brittle
laminae several metres long, L saccharina /
longicruris is suited to extremely sheltered
conditions with little water movement.
Although the three surveys showed a kelp
canopy to be continuously present throughout the
voe, results from transect shrdtes (and also the
ordination analysis) suggest differences in
stability of the vegetation in the inner and outer
vm, The Laminaria hyperborea forests in the
outer areas of Sullom Voe appeared stable in
contrast to the changes in extent and abundance of
Laminaria saccharina / longicruris canopy in the
Sullom Voe suggest that some kelp communities
change more rapidly than others, confirming
WALKER& RICHARDSON'S
(1955) observation
inner voe. However, in the outer voe where that L sacchcrrrna is more noticeably seasonal in
mobile sand is temporarily washed away from its growch than is L hyperborea.
bedrock, the vegetation on uncovered surfaces is
Laminaria I~yperborea also provides habitat
characterised by opportunists such as Saccorhiza for a dynamically stable subflora of epiphytes.
polysch~des and L saccharina / longicruris. The associated underflora assemblage on bedrock
Changes recorded in the inner voe are probably comprised mainly slow-grow~ngspecies that also
natural and possibly cyclical. WALKER
(1 956a) formed a dynamically stable facies. The subtidal
identified changes in the quantity and composition algal underflora communities, and those at
of Laminariaceae in Scotland and showed an 1 1 locations in the v o t without a kelp canopy (see
year cycle and coincidence with sunspot activity; T ~ L E etY a[. 1977 for a full description of these
WALKER(1956b) suggested that this activity may communities), in general agree with the biotopes
influence meteorologica1 conditions and thus the defined by CONNORet al. (1977) from the north
marine environment. Elsewhere in the North of Scotland.
Atlantic grazing by sea-urchins has significantly
The 175 species of algae recorded from
reduced the extent of Laminnria stands ( T r r r ~ v Sullorn Voe (Appendix 1) compares with I99
el al. 1989); this is not the case in Sullom Ym as horn the much longer Hardangerfjord in adjacent
urchins do nut occur in numbers large enough to Norway (JORDE & KLAVESTAD 19631, but
deplete the kelp assemblage. To speculate, the considerably exceeds 49 from SkllaFjadur
contraction of vegetation from deeper waters (comparable in length to Sullom Voe) in the
recorded in the present survey may relate to water Faroes ( T ~ E etYal. 1982).
quality and the flushing of the inner basin of
These subtidal vegetation surveys, and other
Sullom Voe. Although a de-oxygenation event studies over the past twenty years (DUNNET
&
was tecorded in 1978 at the head of Sullom Voe M C T ~ R 199S),
E
have indicated that physical
(PEARSON& ELEFTHER!OU
198 1: STANLEYet al. disturbance caused by construction of the oil
198 I), adverse conditions occurred only beneath a termina1 has been relatively confined. As shown
summer thermocline at 30-35m.
by Walker's many studies LQIninaria populations
The age of the Laminaria hyperborea are resilient to disturbance; h m i n a r i a
population in the outer voe (5-6-7 years) was hyperborea at the oil terminal had recovered in
slightIy more than the 4-5 years reported by 1993 after an absence in 1983.
Walker & Richardson (1957) for open sea
The Laminaria forests that occur on the
populations in Scotland. Although turnover of temperate sea-shores of h e world are biologically
individual L hyperborea plants has occurred diverse and as a consequence are of considerable
between surveys, recruitment has been suFficient conservation impottance. They are described by
to maintain a broadly similar outward appearance BTRKETT
et al. (1998) as the marine equivalent of
of the kelp bed. WALKER
(19581 observed that tropical rainforests in terns of their biological
over a year the standing crop of L hyperboma is productivity, the numbers of individuals 4 i ving
generally stable until reduced by rough seas and there or dependent on the habitat, and the species
by the shedding of old laminae in spring. WALKER diversity present in the habitat. Despite the
(1958) also recorded 'perennial changes' in cover extensive body of knowledge on kelp forests there
and density in Pnminaria populations in which are few long-term studies in Britain. Unlike the
reductions and gains were noted. BIRKE~Tet al. intertidal monitoring in Sullom Voe that was
(1 998) discussed the constant change in the undertaken at shorter intervals and therefore
spatial mosaic of kelp communities and suggested provides more complete time-series information
time-scales of decades to centuries for such on algal populations and communities, these
natural dynamic vanation. Our observations in subtidal algal surveys only provide snapshots at
longer time intervals between which other
unrecorded changes may have occurred.
ACKNOWLEDGEMENTS
We thank Ronnie Gallagher and BP Exploration
Operating Company Ltd., Sullorn Voe Oil
Terminal for the many facilities placed at our
disposal and for considerable support. Shetland
Oil Terminal Environmental Advisoy Group is
gratefully acknowledged for financial support.
The Fieldwork could not have been compfeted
without the enthusrastic participat~on of Lin
Baldock, Jenny Bryant, Frances Dipper. Sarah
Fowler, Rob Huxley, Kay Ingleton, lanny
Maif inson, Chris Spusrier, Karen Webb and Chris
Wood.
REFERENCES
DUNNET.
G.M, & A.D. MCINTYRE.
1995. Monitoring at
an oil terrnlnal: the ShetIand experience.
Proceedir~gs 01ihc Rnyal S o c i ~ f yo f Edinbrirxh
1038: vii + 1-258.
EDMONSTON. A. 1809. A 11icri- of !he a~lcienr and
present S ~ U I Pof ~ l l eShelland Is!and.~.Edlnhurgh
and I a n d ~ nu. i ~+ 345 pp.
GUIRY,M.D. 1997. Benthic, red, brown and green
algae. Pp.341-367 in: HOWSON.C.M. & B.E.
PICTON{Eds). The species directov of t l r ~marine
Jarrr~a and j b r n oJ Ihc Rrirish Isles and
s~irtormdit~g
seas. Ulster Muscum and The Marine
Conservation Saclety, Relfmt and Ross-on-Wye.
vi
TRVINE,
+ 508 pp.
D.E G. 1962. Thr marine algae of Shetland; a
phytogeographical account. British Pl~ycological
%~lle1111
2. 181- 182.
IKVINF,0.E.G 1974. The marine vcgtation of the
Shelland Isle%.Pp. 107-1 13 in: G o o ~ l r R
~ .(Eds).
Tlie natrtroi erlvironm~nroJ Shetland. Proreedi~us
oJ the Nature Consrnrarlcy Council Svmposiult~
held in Edinburgh 29-30 Jarruary 1974.
Edinburgh IV + 178 pp
IRVINE.
D.E.G. 1980. Marine algae. [pp.267-2721. In
BERRY. R J. % 1.L JOHNSTON
(Eds). The Na'atural
Mistor?,of Shetland. Collins. London. 380 pp.
B a r n , D.A.. C.A. MAGGS,
M.J. DRING& P.J.S.
BOADEN.1998. "Infralittoral reer biotopes with
help species" An overview of dynamrc and
sensitivity
characlerrstics for consetvatton IRVINE,D.E.G., M.D. GUIRY. 1. TITTLEY.& G.
RUSSELL. 1975. New and interesting marlne algae
management of m i n e SACS. A r e p e for ha
From the Shelland Isles. Brirish PhycoIogicat
Scorrish Associutiot~ of Marirle Scier~ce. Qt~eetrs
Jortrnal 10: 57-7 I.
University of Belfafl, Porraferry. 174 pp.
B ~ R G W E NF.
, 1903a. Marine algae. Pp.339-532 In: JORDE, I. & N. KUVESTAD. 1963. The Natural History
o f h e F4ardangcrfjord 4, Thc benthonic algal
Warming, E. (Fds). Ro~anyof #he E'aerors based
vegetation. Sarsia 9. 1-99.
upon Danish inv~siigatror~s.
Purr II. Copenhagen.
KAIN, I.M. 1963. Aspects of the b~ologyof Lat~inaricl
681 pp.
I~perborea11. Age, weight, and length. Jonnlal of
BCIRGESEN,F. 1903b. The marine algae of the
I ~ Marrne
P
Rioio~iculAssociatiorl of ihe Utzifed
Shetlnnds. Jotcnlnl of Bofarty, London 41 : 300-386.
Kinxdont 43: 129- 151
BORIRCESEN.
F. & H.J~NSSDN.1905. The distribuhon of
KFN'T, M. & P. COKER.1992. Ve,gelu~iot~
drsrriplion
t h t marine algae of the Arctic Sea and the
and a~lalysisA prac~irol approach. Bcllhaven
northernmost part of thc Atlantic Ocean. Appendix.
Press, London, x +363 pp.
Pp.1-XXVIII. in:WARMING.
E. (Eds). Botarlv oJ'lhe
MDORE, J., P. T A ~ &RK. HISMCK. 1995. Rocky
Faeroes b m d 1tpor1 Dmtish investigarior~s.Parts I .
shores monitoring programme. Proc~edit~gs
of the
2 and 3. Copenhagen. Christiania and London.
Hoyc~lSociet.~of Edirlburpk 103'B. 181-200.
1070 + XXVIlI pp.
.
Ttie benthic
CON?~OR,D.W., M.J D A M N T . 0 H I L ~R.H.F.
.,
HOLT& PEARSON, T.H.& A . E L ~ R I D U1981.
ccolagy of Sullom Voe. Proceedirt~sof ;l~cRo!-aI
W.G.
SANOFBSON.
1997. Marine Nature
Soci~tyof Edirtburgh 8OB:241 -269.
Conservation Rewew: marine hiotope d~sifrcation
SOUTH G.R. & 1. TIITLEY.
1986. A checklist und
for Britain and IreIand. Volume 2. Subltttoral
distribttrianal inde.~01rhe benthic tltarine algae of
biotopes. Version 97.06. JNCC Reporr No. 230: 1rhe N o r ~ h ArIa~tlic Ocean. Huntsman marine
448.
hboratory and British Museum (Narural History).
DLXON, P.S. I963. T h e marine algae of Slietland
St Andrcws and Lnndon. 76 pp
collected during the meeting of the Bntish
STANLEY. S.O., E.E. GRANTHAM,
3 W . LOFTLEY
& N.
Phycological Soc~ety, August 1962. Rrirish
ROBERTSON,N. 1981 Some aspects of sediment
Phycological Ralletir~2: 236-243.
chemistry of SuIIorn Voe, Shetland. Proc~edirlgs
of 1keRoya1Societ,~of Edinburgh SOB: 7 1- !00.
TITLET, I. & W.F. FARNHAM.1997. The srrbtidnl
marine algal vegetation of 5uffo111
Voe reassessed 1994. A report to SOTEAG, The Natural H~story
Museum, London. 155 pp.
%El',
L, W.F. FARNIFAM,
R.L. PLETCIIER
& D.E.G.
IRVME. 1985, The subtidal marine algae of Sullom
Voe reassessed. Transacrions of rht Borat~ical
Soci~tyoJ Edinburgh 44:335-346.
T~LEY
I., , W.F. FARKHAM
& P.W.G GRAY. 19g2.
Seaweeds of the Faroes 2: Sheltered fjords and
sounds. Bullerin oJ ~ i t eBrirish Mtrsetrtr~ (hraruml
History) Botany Serics 10: 133-1 5 1.
T m . I., W.F. FARMIAM.R.G. HooPEu & G.R.
SOUTH.1989. Sublittoral seaweed assemblages 2: a
aansatlantic comparison. Progress in Undemat~r
Scsence 13: 185-20s.
T m w . I., D.E.G. IRVTNE
& N.A. JWHSEN. 1937. The
infralittoral marine algae of Sullo~nVw, Shetland.
Transm~ionsoffhe Botorticnl S o c i e ~of Edirrbtcrgh
42: 397-41 9.
WALKERF.T. t 956a. PeFidicity of the Laminar~aceae
around Scotland. Narure 177: 1246.
WALKERF.T. 1956b. The Lornin~ria cycle. Revue
Algologique 3: 179- 181.
WALKERF.T. 1957. An ecological study of the
Laminariaceae of kilsa Craig. Holy rsland,
Inchmamock, May Island and Seaforth Island.
Tmnsactions ofthe Botanical S o c i ~ ~
ofyEdittbrrrgh
37: 182-199.
WALKER F.T. 1958. Some ecological Factors
conditioning the growth of the Laminariaceate
around Scotland. Actu Adriatira 8 (13): 1-13.
WAL.KFX F T.& W.D, RICHARDSON. 1955. An
ecological rnvcstigation of Ianrinaria cloustonii
h i m . (L hvperborea Fosl.) around Scotland
Journal of Ecology 43: 26-38.
WALKER F.T.& W.D. RICHARDSON1957. Perennial
changes of Larnrrlaria cIous~onSiMrn on the coasts
of Scotland. Journal du Conseil international polir
L'F?.plnralion de la Mer 22: 298-308.
APPENDIX 1: SPECES RECORDED FROM SHETLAND AND SULLOM VOE
Nomenclature and citation mostly follows GUIRY( 1997)
* = Voucher specimens at BM
7 = Recorded from Sullom Voe in 1973
8 = Recorded from Suliorn Voe in 1983
9 = Recorded from SulIom Voe in 1993
Species without asterisks are not represented by specimens at EM; species without 7,8,9 have been
recorded for Shetland but not Sullom Voe.
Bangiophyceae W d o p h y t a )
Aglaothanulion bipinnatum (PCrouan & H Croumj Feldman-Mamyer
dglaolhamnion hookeri (DilFwyn) Magg & Hommmeand)
Aglaolhamnion pricwnum Maggs. Guiry & Rueness
Aglaothamnion roseurn moth) M a g s & L'hatdy-Halos
Aglaorhamnion sepositurrr (Gunnerus) Maggs & Hammersand
Ahnfelt~aplicuta (Hudson) Fries
Aniirhamnio~~~o/loccosa
(0F Muller) Whittick
Asparu~opsisa m t a Harvey [Falkenbergia stage]
Audoui~lcEliaaabrioe (Jbnsson) Woclkerl~ng
Audouinclla h v i e s i i (Dillwyn) Woelkerling
Audouinella mernbmnacea (Magnus) ~ a ~ e n f u s s
Audouinella parvula (KyIm) Dixon
Audouinella purpurea (L~ghtfoot)Woelkerl ing
Audouinella s~cundata(Lyngbye) Dtxon
Audouinella virgatuIa (Harvey) Dixon)
Bangfa atropnrpurea (Roth} C Agardh
Boergeseni~lla
frurict~Iosa(Wulfen) Kylin
Bonnemaisonia h $ e r u Hariot [Trailliella phase]
Brongrziartella byssoides (Godenough & Woodwrtrd) Schmitz
Caldirhamnion c o y t h u m (J E Srmth) Lyngbye
Callithamnion granularurn (Ducluzeau) C Agardh
Cdlirhamnion rerragonum (Withering)S F Gray
Callocolur ngleclus Batters
Callophyllis crisrala (Turner) KUtzing
Callophyllis Eacinida (Hudson) Kiitzing
Catenella caespitosa (WIthering) L Irvine
Ceramiurn cimbricum H Petersen
C~ranliurnakslongchampsii Duhy
Ceramium diaphnum (Lightfoot) Roth
C~mmiumnodulos~m(Lightfoot) Ducluzeau
Cemmiuni shut~lewanhianurn(Kuetzing) Rabenhorst
Cerarnium sfriclum sensir Harvey
Cerarocolax hanzii Rosenvinge
Chondrus ctispur Stac khouse
Chorrrocolaxpolysiphoniar Reinsch
Chylocladin wrticillata (Lightfoot) Blidingia
Coccofylus iruncorus (Palla) M Wynne & J Heine
Colacodictyon reticulaturn (Batters) J Feldrnann
CoralIina off~inalisLinnaeus
Cordylecladia rrecta (Grevllle) J .Agardh
Cruoria pellira (Lyngbye) Eries
Cypropleuru mmosa (Hudson) Lily Newton
CyxrocEo~lilrmpurpurewni (Hudson) Batters
Delesst-ria sunguinea (Hudson) Larnouroux
Qilsea carnosu (Schmidel) Kuntze
Drtmontia contoaa (S Gmelin) Ruprecht
Erythrocladia irrtgu1aris Rosenr~nge
Erythmglossum laciniarun flightfoot) Maggs & Hammersand
Evhrotrichia cornea (Dillwyn) J Agardh
FoslieIla sp.
F~wcellariulimlbrica[is (Hudson) Lamouroux
Gelidirlirmrpusillurn (Stackhouse) Le Jolis
Glarosiphonia capil!aris (Hudson)BerkeIey
? Grecilaria gmcilis (Stackhouse) Stcentoft, L Irvine & Famham
Grifithsia comllitwides (Linnaeus) Trcvisan
I~alamchriionligularum (Woodward) Kuetzing
Fhlosacciocalax kjeflmanii S Lund
I-lalurusJosculos~cs(Lightfoot) Ktitzing
Haraldiophylltml bonnemaisonii (KyIin) A Zinova
Helrninrhora divaxicaru (CAgardh) J Agardh
Hcremsiphonia plut~wsa(Ellis) Batters
Hildcnbrandia cxouanii J Agardh
fliid~nbrandiarubra (SomrnerFeldt) Meneghini
Hypogluss~~m
hypoglossoides IStackhouse) F Collins & Hervey
Kallymenia reniformis (Turner) J Agardh
Lithophyllrrm crouanii Foslie
Lilhophyllum incrustans Phillippi
Lirkophyllum orbiculat~lrn(Foslie) Foslie
Lithothamnron glaciate Kjellman
Litho~horwrionsonderi Hauck
Lomenfaria aniculato (Hudson) Lyngbye
Lumenfaria claveIlasa (Turner) Gaillon
Cornenraria orcndensis ((Harvey)Taylor
Masrocarplls stello~us(Stackhouse) Guiry
Meiodiscus spersbPrgenrts (Kjellman) Saunders k McLachlan
MeloGesia mcmbmrmcea (Esper) Lamouroux
Membranop~eraalatn (Hudson) Stackhousc
Nemotion Iwlminihoide~(VelIcy) Batters
Niiopkyll~rnpunctarum (Stackhouse) GrevilIe
Odonthlia denrara (Linnaeus) Lyngbye
Osmundea hybrid7 (DeCandolle) Nam
Osmundea pinnuiijdu (Hudson) Stackhouse
PaImaria palmara (t~nnaeus)Kuntze
P~yssoneliadubyi P Crorran el H . Crouan
Phycodrys rubens (Linnaeus) Batters
Phyllophom crispa (Hudson) D~xon
Pkyllophom pseudoceranoides IS Gmelin) P Newroth & A R A Taylor
Pkjniarolithon cnlcareum (Fallas) Adey & McKfbbin
Phymnialithon laevigatrrm (Foslie) Foslie
Phymarolirhon Iamii (Lemo~ne)Chamberlain
Phymarolirhon l m o m n d t i (Areschoug) Adey
Phy?natolhhonpurprrreum (P Crouan & H Croum) Woelkcrling & L In
Phymatolirhon ienue (Rosenvinge) Duwel & Wegeberg
P[ocamium carrilagimeum (Linnaeus) Dixon
Plumaria p l m s a (Hudson) Kuntze
*
A ~ ~ o p l ~ y l lc~ilkrpae
um
{P Crouan & H Cwuan) P Jones & Woelkerling
Pneophyllurn canfervicala (Kuetzing) Chamberlain
Pubides mrllndits (Hudson) Greville
*
rp
Polysiphonia at furlricaKapraun & J Norris
PoIysiphonia brodiaui (Dillwyn) Sprengel
*
P~l~vsiphonia
~longoro(Hudson} Sprengl
*
Pnlysrphoniu elori~elInHarvcy
PoIysiphoniaJbror~ (Dillwyn) Harwey
IP
PoIvsiphotiia~~riIIosu
(Dillwyn) Sprengel
Polysiphonin Jucaides (Hudson) GreviIle
c
Polyiphonia Iarrosa (Linneaus)Tandy
*
PoIysiphorliu tiigro (Hudson) Batten
s
Polysipl~o~lia
stricra (Dl1l w yn) GreviIle
*
Porpiryra amplissinta (Kjellman) Setchell & Hus
Porplyra leucosricra Thuret
*
Porpkyra umbilicalis (Linnaeus) KUtzing
r~
Porpkyropsis coccirl~a(Areschoug) Rosenvinge
XI
Porpk~rosrsomiumcilia re (Harvey) M Wynne
Plcrosiphoriia pamsirica (Hudson) Edkenberg
*
Pr~rolhanmionplumrtla (Ellis) Nageli
4r
Pfilota gnllneri P Silva, Maggs & L irvine
*
PfiIithaninio~pluma [Dillwyn) Thuret
Rlzadomelo confentoides (Hudson) P Sjlva
Rhodomela iycopodioides (Linnaeus) C Agardh
I
RhodophyIlis divaricara (Stackhouse) Papenfuss
4
v6
Rhuduphyserna elegans (J Agardh) Dixon
Rho$o~lranir~ielia~idula
(Dl1lwyn) J Feldrnann
*
Sahlr~igiasubinregm (Rosenvingc) Kornmann
Scagelia pilsilla (Ruprecht) Athanasladrs
*
Scinuia furceliota (Turner) J kgardh
v
Seirospom intermpta (J E Smith) Schmitz
*
Sp~rnlofhaa~nion
repens (Dillwyn) Rosenvinge
St$onema alsidii (Zanardini) K Drew
Tztanodernro carallinae (P Crouan & H Crouan)Woelkerling, Chamberlain & P Silva
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
7
T
Fucophyceae (Phaeophy ta)
Acineiespara crinito (Harvey) Kornmann
Rcrollrrix gracilis Kylin
Alaria esculenlo (Llnnaeus) Greville
Ascophyllun~tlodosuni (Linnaeus) Le Jolis
Asperococc~lsconlpressrrs W J Hooker
Asperncocct~sJsrulosus(Hudson) W J Hooker
Asperococcr~.~
bidlnsus Larnouroux
Cliilionen~aoccllaftcn~(Kuetzing) Kuckuck
Clrilionrmo mpmns ( P Crouan et H Crouan) Sauvageau
Chordafilun~(Linnaeus) Stackhouse
CkordariaJagell$wrmis (0F Miiller) C Agardh
Cba$ostep h m spongiosus (Hudson) C Agardh
Colpomenia per~grina(Sauvageau) C Agard h
Curleria mulrfida (Menens) De Notaris [includes Aglaozonia stage]
Desmareslia aculeara (L~nnaeus)Lamoumux
D~smaresriraligltlata (Lightfoot) Lamouroux
Desnluresria viridis (0F Miiller) Larnouroux
a
7
*
7
7
*
*
rt-
Dic~yosiphonchordaria Areschoug
Dicryusiphon j o ~ n i c ~ ~ l a c(1-ludson)
~us
Grevil le
Dicryora djchofoma (Hudson) Lamouroux
EcrocurpusfascicuIaf~~s
Harvey
Ecfocarpt~ssrlrcrtloszrs (Dillwyn) Lyngbyc
Etnchis1aj7accida [Dtllwyn) Areschoug
Eiachistafi~cicola(Velicy) Arcschoug
E/uchistra scli/irla:a (J E Smith) Duby
Endodictyorl irlfesiarts Gran
Eudes~rievirescens (Berkeley) J Agardh
Frdcus ceranoides Linnaeu s
Flrcrrs distichus Linnaeus
Fuelis evanescetis C Agardh
F'ucus serrartrs Linnaeus
firctls spirnlis Li nnaeus
Ftrc~tst ~ s i a t l o s ~Linnaeus
rs
Giraridia sphacelnrioid~sDerbts & Solier
Got~ot~enra
aecidioides (Rosenvinge) P Pedersen
Halidqx siliqrtosa (Linnaeus) Lyngbye
Ifecatnncmo r~racttlans(F Collins) Sauvagcau
llerpotienta lvlfi?iniirrt(Greville) iAgardh
Hinienrhalia e l o ) ~ g a (Linnaeus)
~a
S Gray
Hincksia granrdom (1 E Srn~tli)P Silva
Ilirtcksia hincksia~(Harvey) P Silva
Hincksia ovara ( K j d 1 man)P Silva
Hi~icksiascr~idriana(Znanrdini) P Silva
Hincksia secunda (Kuctzing) P Silva
Isrhr~lople~~
~phaerophora(Harvey) KjcIlman
Larrli?iariu drgirora (Hudson) Lamouroux
Lamb~arial~yperborea(Gunnerus) Foslie
Laminnria longicnrris De la Pylaie
Laminaria succharina (Linnaeus) hrnoumux
hminariocohr tome~rrosoides(Farlow) Kylin
barbesiu d v f o m ~ i s(L~nnaeus)Areschoug
Lep?oriemarellafasriarlafu (Reinke) P Silva
Lzfosiphon lar~tit~ariae
(Lyngbye) Rarvcy
Mesogloia lariosa P Crouan & H Crouan
M~sogloiaven~~iutlatu
(J E Srn~th)S Gray
Mikrosypharpo~siphorriueKuckuck
Mikrosyphar porpkymr Kuckuck
Mvriacrula ar~schorrgii(P Croi~an& H Crouan) G Hamel
Myriactttln clandesrina (P Crouan & H Crouan) J Fe ldrnann
Myrionenln corrrnnac Sauvageau
Mvriot~e~rur
papillosum Sauvageau
12.lvrioncrnas~rangrilansGreville
Myriofn'chia cla~wejormisHarvey
Pelveria canalicr~lara(Linnaeus)Decaisne & Thurel
Peralor~iafascia (0F Mliller) Kuntze
P~rrodermamacriliforr~~e
(Wollny) Kuckuck
Phaeostronia ptistulosunl Kuckuck
Pilay~llaiirroralis (I-innacus) KjeZlman
Pogorrich~mlfilifoormp Reinke
Prorecrocarplis specios~ds(R@rgesen)Kommann
Pscrrdolithodennu exrensum (PCmuan & H Crouan) S Lund
Puncia ria larifulia Greveille
Puncraria pIanlaginea (Roth) Grrville
Punclaria tenuissima (C Agardh) Greville
Ralfsia wrrucosa (Areschoug) Areschoug
Saccorhim polysckides (Lightfoot)Batters
Sauvageaugloia chordariaebrmis (P Crouan & H Crouan) Kylin
Sauva~eaugloiagrtJ5rhsiana (W Hooker) Kylin
Scyfosipkon lamentaria (Lyngbye) Link
Sorocarpur micromoms (Bory) P Silva
Spcrmatochnw paradoxus (Roth) Kiltzing
Sphacelaria arcrica Hmey
Sphacelaria cirrosa (Roth)C Agardh
Sphacelaria fusca (Hudson) S Gray
Sphacelaria rnirabilis (Batters) Prud'hommc van Reine
Sphacelaria nana Kiitzlng
Sphacelaria plumig era Aauck
Sphacelaria plrrmosa Ly ngbye
Spkacelnria rigidula Kuetzing
Sphacrotrichia divaricata (CAgardh) Kylin
Spongonema romnrosum (Hudson) Kiltzing
Srlcyrosipkon grifithianus (Lelolis) Helmes & Batters
S~ictyosiphonsclrifems (Reinke) Rosenvinge
d~ictyosiphonfortilis (Ruprecht) Reinke
Stilophora tenella (&per) P Silva
Srmgularia clavara ( H m c y ) G Hamel
Streblonem breve (Sauvageau) De Toni
Streblonemafascicsllaium Thuret
Srreblonema pamsiricum (Sauvageau) Levnng
Streblonerna sphericurn (Derbks L Solier) Thuret
UIonema rhizophom Fodie
Chlorophyceae
Acrochaere leprorhaere (Huber) R Nielsen
Acrockaete viridis (Reinke) R Nielsen
Acrochaete winrockii (Wille) R Nielsen
BLasiophysa rhizoptts Reinke
Blidingim minima (Klitang) Kylin
Bulbocoleon piliferum N Priagsheim
Btyopsis hypnoides Lamouroux
Bryopsis plumosa (Hudson) C Agardh
Capsosiphon fulvesc~r~s
(C Agardh) Setchell L N Gardner
Chaeromotpha linurn (0F Mtiller) KUtzing
Chaeiomurgha medirerranm (KUtz~ng)Kiibing
Chaeromorpka melag~nium(Weber et Mohr) Kiitzing
CIadophora albida (Nees) KUtzing
Cladophom dalmarica Kutzing
Cladophom hutchinsiae (Dillwyn) Kiltzing
Cladophora pygmaea Reinke
Clodophom rupestris (Linnaeus) KIitzing
Cladophm sericea (Hudson) KUtzing
Codiumfragile (Suringar) Hariot ssp. atlanticurn {Cotton) P Silva
Derbesia marina (Lyngbye) Solier [inctud~ngHalicystis stage]
Enterornorpha clnthro~a(Roth) Greville
Enr~romorphacompressa (Linnaeus) Nees
Enteromorpha flexuasa (Wulfen) J Agardh
Enteromorpha intestinalis (Linneaus) Nees
Enleromorpha muscoides (Clemente y Rubio) Cremades
Enieromorpha prolifera (0F MIiller) J Agardh
E~eromorphararsii Harvey
Entocladia pejorans (Huber) Levring
EpicludiaJustmt Reinke
Eugomontia saccula~aKomrnann
Ochlochaetejerox Thwaites
Ostroebium queketii Bornet B! Flahault
Percursaria percursa (C Agardh) Rosenvinge
Pmsiola crispa (Lightfoot) Kiiking
Prasiola stipitala Jessen
Pkaeophila dendroides (PCrouan & H Crouan) Batters
Pringsheirniclla scutata [Reinke) Marchewianka
Pscudodicryon injlalum Ercegovic
Psedendocfoniuni fucicola (Rosenvinge) R Nielsen
Rhizoclonim turtuosum (Diilwyn) Kutzing
Spongomorpha arcfa (Dillwyn) Kiltzing
Spongonenla acmginosa (Linnaeus) van den Boek [Chlomhytrium inclusum stage]
Ulo~hrixJacca(Dl llwyn) Thuret
Ulothrix speciosa (Harvey)Kiltzing
*
Ulva lacruca Linnaeus
Ulva rig ida C Agardh
Ulvaria juscra (Pastels & Rupre~ht)Ruprecht
UIveila !ens P Crouan & H Crouan
UEvopsis grevillei (Thuret) Gayral
Ilrospora penicilliforms (Roth) Areschoug