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New records of benthic marine algae for Norway, with notes on some rare species 

from the Florø district, western Norway 

Tor Eiliv Lein, Grethe Bruntse, Karl Gunnarsson & Ruth Nielsen 

SARSIA 

INTRODUCTION 

Lein TE, Bruntse G, Gunnarsson K, Nielsen R. 1999. New records of benthic marine algae for Norway, 

with notes on some rare species from the Florø district, western Norway. Sarsia 84:39-53. 

An investigation of the algal vegetation in the Florø district was undertaken in 1997. Representative 

samples were selectively collected at exposed and sheltered sites in February and September. New to 

Norway were: Acrochaete heteroclada, A. inflata, Aglaothamnion cf. pseudobyssoides, and Ulvella 

setchellii. Ceramium botryocarpum, C. pallidum and C. siliquosum have not previously been recorded 

in Norway, but may have been reported under different synonyms. Epicladia philipsii, Gracilariopsis 

longissima, Pseudendoclonium dynamenae, and Syncoryne reinkei were recently separated from other 

species and old records of these may in part cover the new species. The study extends the known 

northern boundary of distribution along the Norwegian west coast for 20 species. Reproductive male 

plants of Rhodochorton purpureum were observed, probably for the first time in nature. A summary of 

all species known to occur in the district is given and includes 33 Nostocophyceae, 121 Bangiophyceae, 

64 Fucophyceae, and 56 Chlorophyceae. The distribution of some species in certain west Norwegian 

districts are discussed in relation to relatively warm winter temperatures. 

Tor Eiliv Lein, University of Bergen, Department of Fisheries and Marine Biology, PO Box 7800, 

N-5020 Bergen, Norway. – Grethe Bruntse, Kaldbak Marine Biological Laboratory, FR-180 Kaldbak, 

Faroe Islands. – Karl Gunnarsson, Marine Research Institute, 121 Reykjavik, Iceland. – Ruth Nielsen, 

Botanical Museum and Library, University of Copenhagen, DK-1123 Copenhagen K, Denmark. 

E-mail: tor.eiliv.lein@ifm.uib.no – gbruntse@hotmail.com – karl@hafro.is – ruthn@bot.ku.dk 

Keywords: Benthic algae; Nostocophyceae (Cyanophyceae); Bangiophyceae; (Rhodophyceae); 

Fucophyceae (Phaeophyceae); Chlorophyceae; Biogeography; Sogn og Fjordane; Norway. 

The marine algae of western Norway are relatively well known 

(Kylin 1910; Printz 1926; Hygen & Jorde 1935; Levring 

1937; Jorde & Klavestad 1963). However, a recent compilation 

of the distribution of marine benthic macroorganisms in 

26 sectors along the Norwegian coast (Brattegard & Holthe 

1997) made it clear that the algal vegetation is poorly known 

in some regions, particularly the northern part of Sogn og 

Fjordane county (sector 10, Fig. 1). 

Indeed, the only published records of marine benthic 

algae for this area are a thesis based on material collected 

in August 1994 (Brattenborg 1997), and a 

preliminary list of species from the same survey (Lein 

& al. 1994), which also includes older records collected 

at the end of the 19th century and maintained at the 

Botanical Museum, University of Bergen (BG). 

An investigation to the south of the area (sector 9, 

Boye 1896), indicated a rich marine vegetation including 

many rare species known only from here or a few 

other sites in Norway. Reinvestigation in 1994-96 of 

the vegetation at the sites visited by Boye 100 years 

ago confirmed the presence of rare and unusual algae 

for Norway (Lein 1996). 

This paper contributes to the knowledge of the distribution 

of benthic algae in western Norway with 

records of new and rare species observed in the Florø 

district, 1997. The paper also presents an updated list 

of all marine algae known from this area, sector 10 in 

Brattegard & Holthe (1997). 

AREA DESCRIPTION 

Collections were made in the outer coastal area in the 

Florø district, western Norway in February and September 

1997 at 44 stations located in 17 sampling areas 

(Fig. 1). The area is characterised by numerous islands 

and skerries and highly indented coastlines. 

Wave exposure of the stations varied from exposed to 

sheltered, and was estimated according to Baardseth 

(1970), see Lein & al. (1998). Tidal currents were probably 

significant at some stations, but no data are available. 

The mean monthly temperatures (°C) and salinity values 

at 4 m depth for the period 1936-1970 are available 

for Stad, a site at the northern border of sector 10 

(Midttun 1975). Mean temperature for the period 1936- 

1970 was 4.6 °C (SD = 1.0) in March and 13.9 °C (SD = 

2.5) in August, the coldest and warmest month respec-

40 Sarsia 84:39-53 – 1999 

Fig. 1. Location of sampling areas. Insert shows map of Norway with the sectors 1-26 according to Brattegard & Holthe (1997). 

tively. The mean monthly salinity values for the same 

period were 32.4-33.2 psu from December to June and 

30.8-31.7 psu from July to November. In 1997 the mean 

monthly temperature and salinity values were 5.2 °C and 

32.9 psu for March and 17.6 °C and 29.1 psu for August 

(data from the Institute of Marine Research, Bergen). 

Temperature and salinity values at hydrographic stations 

in the Florø district were measured during the sampling 

periods and corresponded well with the mean temperature 

and salinity for Stad, except in two areas that 

had a fresh water supply from small local streams, see 

Lein & al. (1998). 

MATERIAL AND METHODS 

In February the material was collected by dredge in the 

sublittoral zone, except for two samples obtained by 

skin diving in the sublittoral fringe. In September the 

samples were hand picked in the littoral zone and ob- 

tained by skin- or scuba diving in the sublittoral zone 

to a depth of 30 m. Detailed information on sampling 

area, positions (WGS-84), depth (m), date and method 

used at each station are given in Lein & al. (1998). 

Most of the samples obtained in February were fixed 

in 4 % formaldehyde in sea water immediately after collection 

and later rinsed in fresh water before being studied. 

In September the samples were examined while fresh 

at the Marine Biological Station, Espegrend. Microphotographs 

of KARO embedded specimens were obtained 

using Olympus AX 70 microscope equipped with 

Normarski differential interference contrast optics (DIC). 

To confirm the identity of some of the small epiphytic 

green algae, cultures were established from fragments 

of host-plants with epiphytes. The cultures were kept 

in Petri dishes with MV 30 as culture medium 

(Christensen 1982), initially 5 mg l –1 of GeO 2 was added 

to suppress growth of diatoms. The dishes were incubated 

at 15 °C with an irradiance of 20 µmol m –2 s –1

photon flux density on a 16:8 h light:dark cycle. 

The Corallinales were dried and decalcified in 0.6 M 

H 2 NO 3 before further treatment. The material was treated 

as described by Düwel & Wegeberg (1996) or embedded 

in Cryo-Embed (AX-LAB A/S, Copenhagen, Denmark) 

to obtain sections of 7-10µm with a Reichert-Jung 2800 

Frigocut freeze microtome. The sections were coloured 

with 10 % aqueous Cotton Blue and mounted in KARO. 

Permanent slides and herbarium material have been 

deposited in the Botanical Museum, University of 

Bergen (BG) and at the Botanical Museum, University 

of Copenhagen (C). Syntype of Lithothamnion 

norvegicum was obtained from the Rijksherbarium, 

University of Leiden (L). 

The nomenclature generally follows Nielsen & al. 

(1995). The Nostocophyceae are named according to 

Anagnostidis & Komarek (1985, 1988, 1990), 

Anagnostidis & Pantazidou (1991) and Komarek & 

Anagnostidis (1986, 1989), while the Corallinales follow 

Irvine & Chamberlain (1994). 

RESULTS 

The substrate at almost all stations were dominated by 

bedrock or rocks with patches of shellsand and fine 

sediments in sheltered areas. At some stations sediments 

composed of small stones, pebbles, shellsand or mud 

were dominant. Laminaria hyperborea or L. saccharina 

were the main vegetations on most sublittoral hard bottom 

stations depending on wave exposure. The two most 

sheltered freshwater influenced stations, featured rock 

covered by crustose algae and low growing red algae, 

and mud covered with Zostera marina L. respectively; 

Table 1. Recorded Nostocophyceae (Cyanophyceae). 

Anabaena torulosa 

(Carmich. ex Harv. in Hook.) Bornet & Flahault 

Aphanothece pallida (Kütz.) Rabenh. 

Arthrospira major (Kütz.) Crow 

Calothrix aeruginea (Kütz.) Thur. ex Bornet & Flahault 

Calothrix confervicola (Roth) C. Agardh ex Bornet & Flahault 

Calothrix scopulorum 

(F. Weber & D. Mohr) C. Agardh ex Bornet & Flahault 

Chroococcus dimidiatus (Kütz.) Nägeli 

Dermocarpa acervatus (Setch. & N.L. Gardner) Pham-Hoang Ho 

Dermocarpa kerneri (Hansg.) Hansg. 

Dermocarpa schousboei 

(Thur. in Bornet & Thur.) Bornet in Batters 

Gloeocapsopsis crepidinum (Thur.) Geitler 

Heteroleibleinia infixa (Frémy) Anagnost. & Komarek 

Leibleinia epiphytica (Hieron.) Anagnost. & Komarek 

Leibleinia willei (Setch. & N.L. Gardner) P.C. Silva 

Leptolyngbya nostocorum 

(Bornet ex Gomont) Anagnost. & Komarek 

Leptolyngbya terebrans (Bornet & Flahault) Anagnost. & Komarek 

Lein & al. – New records of benthic algae for Norway 41 

see Lein & al. (1998) for more details. 

The observed species of Nostocophyceae, 

Bangiophyceae, Phaeophyceae, and Chlorophyceae together 

with other records from the area appear in Tables 

1 and 2. Included are species reported for sector 

10 by Rueness & al. (1997a, b, c), verified algae observed 

by Brattenborg (1997), and unpublished data 

from student courses in October 1996 and April 1997 

(see Lein & al. 1998). 

Notes are given for species that are new to Norway 

(indicated by § or (§)), southern species for which the 

record extends the known northern boundary of distribution 

along the Norwegian coast (#) and for species 

that are considered rare, having been recorded by 

Rueness & al. (1997a, b, c) in three or fewer sectors or 

not at all. Also included are notes on species with special 

reproductive structures or taxonomic problems. 

NOSTOCOPHYCEAE (CYANOPHYCEAE) 

In most of our collections several species of blue-green 

algae were recorded and a preliminary list is presented 

as Table 1. 

The taxonomic problems with delimitation of species 

within this group demand a thorough and critical 

review of published records before it is possible to compile 

a complete, commented species list for the areas, 

or a distributional table for the Nostocophyceae of Norway. 

Furthermore, published records in recent publications 

are scarce (Munda 1967; Wiik 1981; Molvær & 

al. 1984; Pedersen & al. 1985), and this group of organisms 

was not included among the tables by Brattegard 

& Holthe (1997). 

Lyngbya confervoides (C. Agardh ex Gomont) Gomont 

Lyngbya semiplena (C. Agardh) J. Agardh 

Mastigocoleus testarum Lagerh. 

Microchaete grisea Thur. ex Bornet & Flahault 

Microcystis litoralis (Hansg. in Foslie) Forti in De Toni 

Nostoc entophytum Bornet & Flahault 

Oscillatoria limosa C. Agardh ex Gomont 

Phormidium chalybeum 

(Mert. ex Gomont) Anagnost. & Komarek 

Phormidium corallinae (Kütz.) Anagnost. & Komarek 

Porphyrosiphon martensianus 

(Menegh. ex Gomont) Anagnost. & Komarek 

Rivularia atra Roth ex Bornet & Flahault 

Schizothrix tenerrima (Gomont) Drouet 

Sphaenosiphon olivaceus Reinsch 

Sphaenosiphon prasinus Reinsch 

Spirulina subsalsa Oerst. ex Gomont 

Stanieria sublitoralis (Lindstedt) Anagnostidis & Pantazidou 

Tolypothrix tenuis Kütz.

42 Sarsia 84:39-53 – 1999 

Table 2. Bangiophyceae (Rhodophyceae), Fucophyceae (Phaeophyceae), and Chlorophyceae known for sector 10. § New record 

for Norway. (§) Record of a recently established species, possibly recorded under a different name in older publications. # Record 

extends known northern boundary of distribution in Norway. 

Bangiophyceae (Rhodophyceae) 

Acrochaetium collopodum (Rosenv.) Hamel 

Acrochaetium parvulum (Kylin) Hoyt 

# Acrochaetium reductum (Rosenv.) Hamel 

Acrochaetium secundatum (Lyngb.) Nägeli 

Aglaothamnion bipinnatum 

(P. Crouan & H. Crouan) Feldm.-Maz. 

Aglaothamnion hookeri (Dillwyn) Maggs & Hommers. 

§# Aglaothamnion cf. pseudobyssoides 

(P. Crouan & H. Crouan) Halos 

Aglaothamnion scopulorum 

(C. Agardh) Feldmann-Mazoyer 

Aglaothamnion sepositum 

(Gunnerus) Maggs & Hommers. 

Aglaothamnion tenuissimum (Bonnem.) Feldm.-Maz. 

Ahnfeltia plicata (Huds.) Fr. 

Antithamnionella floccosa (O.F. Müll.) Whittick 

Apoglossum ruscifolium (Turner) J. Agardh 

Audouinella efflorescens (J. Agardh) Papenf. 

Audouinella membranacea (Magnus) Papenf. 

Audouinella pectinata (Kylin) Papenf. 

Bangia atropurpurea (Roth) C. Agardh 

Bonnemaisonia asparagoïdes (Woodw.) C. Agardh 

Bonnemaisonia hamifera Har. 

Brongniartella byssoïdes (Gooden. & Woodw.) F. Schmitz 

Callithamnion corymbosum (Sm.) Lyngb. 

Callithamnion tetragonum (With.) Gray 

# Callocolax neglectus Schmitz ex Batters 

Callophyllis cristata (C. Agardh) Kütz. 

Callophyllis laciniata (Huds.) Kütz. 

(§)# Ceramium botryocarpum J.W. Griff. ex Harv. 

Ceramium cimbricum H.E. Petersen in Rosenv. 

Ceramium deslongchampii Chauv. ex Duby 

# Ceramium diaphanum (Lightf.) Roth 

Ceramium nodulosum (Lightf.) Ducluz. 

(§) Ceramium pallidum 

(Nägeli ex Kütz.) Maggs & Hommersand 

Ceramium secundatum Lyngb. 

Ceramium shuttleworthianum (Kütz.) Rabenh. 

(§) Ceramium siliquosum (Kütz.) Maggs & Hommers. 

Ceratocolax hartzii Rosenv. 

Chondrus crispus Stackh. 

Choreocolax polysiphoniae Reinsch 

Chroodactylon ornatum (C. Agardh) Basson 

Chylocladia verticillata (Lightf.) Bliding 

Coccotylus truncatus (Pall.) M.J. Wynne & J.N. Heine 

Colaconema daviesii (Dillwyn) Stegenga 

# Colaconema nemalii (De Not. ex L. Dufour) Stegenga 

Colaconema savianum (Menegh.) R. Nielsen 

# Colaconema strictum (Rosenv.) R. Nielsen 

# Compsothamnion thuyoides (J.E. Smith) Nägeli 

Corallina officinalis L. 

Cruoria pellita (Lyngb.) Fr. 

Cryptopleura ramosa (Huds.) Kylin ex L. Newton 

Cystoclonium purpureum (Huds.) Batters 

Delesseria sanguinea (Huds.) J.V. Lamour. 

Dilsea carnosa (Schmidel) Kuntze 

Dumontia contorta (S.G. Gmel.) Rupr. 

Erythrocladia irregularis Rosenv. 

Erythrodermis traillii (Holmes ex Batters) Guiry & Garbary 

Erythrotrichia carnea (Dillwyn) J. Agardh 

Erythrotrichia reflexa 

(P. Crouan & H. Crouan) De Toni sensu Rosenv. 

Furcellaria lumbricalis (Huds.) J.V. Lamour. 

Gelidium latifolium (Grev.) Bornet & Thur. 

Gloiosiphonia capillaris (Huds.) Carmich. ex Berk. 

Gracilaria gracilis 

(Stackh.) M. Steentoft, L.M. Irvine & W. Farnham 

(§) Gracilariopsis cf. longissima (S.G.Gmelin) 

M. Steentoft, L.M. Irvine & W. Farnham 

Griffithsia corallinoides (L.) Trevis. 

Haemescharia hennedyi 

(Harv.) K.L. Vinogr. & T.A. Jacovleva 

# Halurus flosculosus (J.Ellis) Maggs & Hommersand 

Heterosiphonia plumosa (J.Ellis) Batters 

Hildenbrandia rubra (Sommerf.) Menegh. 

Lithophyllum crouanii Foslie 

Lithothamnion corallioides 

(P. Crouan & H. Crouan) P. Crouan & H. Crouan 

Lithothamnion glaciale Kjellm. 

Lithothamnion norwegicum (Aresch.) Kjellm. 

Lithothamnion sonderi Hauck 

Lomentaria clavellosa (Turner) Gaillon 

Lomentaria orcadensis (Harv.) Collins ex W.R. Taylor 

Mastocarpus stellatus 

(Stackh. in With.) Guiry in Guiry et al. 

Meiodiscus spetsbergensis 

(Kjellm.) G.W. Saunders & McLachlan 

Melobesia membranacea (Esper) J.V. Lamour. 

Membranoptera alata (Huds.) Stackh. 

Monosporus pedicellatus (Sm.) Solier 

Nemalion multifidum (F. Weber & D. Mohr) Endl. 

Nitophyllum punctatum (Stackh.) Grev. 

Odonthalia dentata (L.) Lyngb. 

? Osmundea truncata (Kütz.) Nam & Maggs 

Palmaria palmata (L.) Kuntze 

Peyssonnelia dubyi P. Crouan & H. Crouan 

Phycodrys rubens (L.) Batters 

Phyllophora crispa (Huds.) P.S. Dixon 

Phyllophora pseudoceranoïdes 

(S.G. Gmel.) Newroth & A.R.A. Taylor 

Phymatolithon lenormandii (Aresch.) W.H. Adey 

Phymatolithon purpureum 

(P. Crouan & H. Crouan) Woelk. & L.M. Irvine 

Phymatolithon tenue (Rosenv.) Düwel & Wegeberg 

# Plagiospora gracilis Kuck. 

# Pleonosporium borreri (Sm.) Nägeli 

Plocamium cartilagineum (L.) P.S. Dixon 

Plumaria plumosa (Huds.) Kuntze 

Pneophyllum fragile Kütz. 

Polyïdes rotundus (Huds.) Gaillon 

Polysiphonia brodiaei (Dillwyn) Spreng.

Table 2. Continued. 

Polysiphonia elongata (Huds.) Spreng. 

Polysiphonia fibrillosa (Dillwyn) Spreng. 

Polysiphonia fucoides (Huds.) Grev. 

Polysiphonia lanosa (L.) Tandy 

Polysiphonia stricta (Dillwyn) Grev. 

Porphyra miniata (C. Agardh) C. Agardh 

Porphyra purpurea (Roth) C. Agardh 

Porphyra umbilicalis (L.) J. Agardh 

Porphyropsis coccinea (J. Agardh ex Aresch.) Rosenv. 

Pterosiphonia parasitica (Huds.) Falkenberg 

Pterothamnion plumula (J. Ellis) Nägeli 

Ptilota gunneri P.C. Silva, Maggs & L.M. Irvine 

Rhodochorton purpureum (Lightf.) Rosenv. 

Rhodomela confervoides (Huds.) P.C. Silva 

Rhodomela lycopodioides (L.) C. Agardh 

Rhodophyllis divaricata (Stackh.) Papenf. 

Rhodophysema elegans 

(P. Crouan & H. Crouan ex J. Agardh) P.S. Dixon 

Sahlingia subintegra (Rosenv.) Kornmann 

Scagelothamnion pusillum (Rupr.) Athanas. 

# Schmitzia hiscockiana Maggs & Guiry 

Seirospora interrupta (Sm.) F. Schmitz 

Spermothamnion repens (Dillwyn) Rosenv. 

Stylonema alsidii (Zanardini) K.M. Drew 

# Titanoderma laminariae 

(P. Crouan & H. Crouan) Y.M. Chamb. 

Titanoderma pustulatum (J.V. Lamour.) Nägeli 

Fucophyceae (Phaeophyceae) 

Acrothrix gracilis Kylin 

Alaria esculenta (L.) Grev. 

Ascophyllum nodosum (L.) LeJol. 

Asperococcus bullosus J.V. Lamour. 

Asperococcus fistulosus (Huds.) Hook. 

Chilionema ocellatum (Kütz.) Sauv. 

Chilionema reptans (P. Crouan & H. Crouan) Sauv. 

Chorda filum (L.) Stackh. 

Chordaria flagelliformis (O.F. Müll.) C. Agardh 

Cladostephus spongiosus (Huds.) C. Agardh 

Colpomenia peregrina (Sauv.) Hamel 

Cutleria multifida (Sm.) Grev. 

Desmarestia aculeata (L.) J.V. Lamour. 

Desmarestia ligulata (Lightf.) J.V. Lamour. 

Desmarestia viridis (O.F. Müll.) J.V. Lamour. 

Dictyosiphon foeniculaceus (Huds.) Grev. 

Dictyota dichotoma (Huds.) J.V. Lamour. 

Ectocarpus fasciculatus Harv. 

Ectocarpus siliculosus (Dillwyn) Lyngb. 

Elachista fucicola (Velley) Aresch. 

Elachista stellaris Aresch. 

Fucus distichus L. 

Fucus serratus L. 

Fucus spiralis L. 

Fucus vesiculosus L. 

Giraudia sphacelarioides Derbès & Solier in Castagne 

Halidrys siliquosa (L.) Lyngb. 

Himanthalia elongata (L.) Gray 

Hincksia hincksiae (Harv.) P.C. Silva 

Hincksia ovata (Kjellm.) P.C. Silva 


Hincksia sandriana (Zanardini) P.C. Silva 

Isthmoplea sphaerophora 

(Carmich. ex Harv. in Hook.) Kjellm. 

Laminaria digitata (Huds.) J.V. Lamour. 

Laminaria hyperborea (Gunnerus) Foslie 

Laminaria saccharina (L.) J.V. Lamour. 

Laminariocolax tomentosoïdes (Farl.) Kylin 

Leathesia difformis (L.) Aresch. 

Leptonematella fasciculata (Reinke) P.C. Silva 

Litosiphon laminariae (Lyngb.) Harv. 

Mesogloia vermiculata (Sm.) Gray 

Mikrosyphar polysiphoniae Kuck. 

# Myriactula chordae (Aresch.) Levring 

Myrionema corunnae Sauv. 

Myrionema foecundum (Strömf.) Sauv. 

Myrionema magnusii (Sauv.) Loiseaux 

Myriotrichia clavaeformis Harv. 

Pelvetia canaliculata (L.) Decne. & Thur. 

Petalonia fascia (O.F.Müll.) Kuntze 

Petalonia zosterifolia (Reinke) Kuntze 

Pilayella littoralis (L.) Kjellm. 

Protectocarpus speciosus (Børgesen) Kornmann in Kuck. 

Pseudolithoderma extensum 

(P. Crouan & H. Crouan) S. Lund 

Ralfsia verrucosa (Aresch.) Aresch. in Fries 

Scytosiphon lomentaria (Lyngb.) Link 

Spermatochnus paradoxus (Roth) Kütz. 

Sphacelaria caespitula Lyngb. 

Sphacelaria cirrosa (Roth) C. Agardh 

Sphacelaria plumosa Lyngb. 

Sphacelaria plumula Zanardini 

Sphacelaria radicans (Dillwyn) C. Agardh 

Sphacelaria rigidula Kütz. 

Spongonema tomentosum (Huds.) Kütz. 

Stictyosiphon soriferus (Reinke) Rosenv. 

Stictyosiphon tortilis (Rupr.) Reinke sensu Rosenv. 

Chlorophyceae 

§# Acrochaete heteroclada J.A. Correa & R. Nielsen 

§# Acrochaete inflata (Erce.) Gallardo et al. 

Acrochaete repens Pringsh. 

Acrochaete viridis (Reinke) R. Nielsen 

Acrochaete wittrockii (Wille) R. Nielsen 

Acrosiphonia arcta (Dillwyn) Gain 

Blastophysa rhizopus Reinke 

Blidingia minima (Nägeli ex Kütz.) Kylin 

Bolbocoleon piliferum Pringsh. 

Bryopsis plumosa (Huds.) C. Agardh 

Chaetomorpha ligustica (Kütz.) Kütz. 

Chaetomorpha linum (O.F. Müll.) Kütz. 

Chaetomorpha melagonium (F. Weber & D. Mohr) Kütz. 

Chlorocystis cohnii (E.P.Wright) Reinhard 

Cladophora albida (Nees) Kütz. 

# Cladophora dalmatica Kütz. 

Cladophora flexuosa (O.F. Müll.) Kütz. 

# Cladophora pygmaea Reinke 

Cladophora rupestris (L.) Kütz. 

Cladophora sericea (Huds.) Kütz. 

Codium fragile (Suringar) Har.

44 Sarsia 84:39-53 – 1999 

Table 2. Continued. 

Derbesia marina (Lyngb.) Solier 

Enteromorpha compressa (L.) Nees 

Enteromorpha flexuosa (Wulfen) J. Agardh 

Enteromorpha intestinalis (L.) Nees 

Enteromorpha linza (L.) J. Agardh 

Enteromorpha prolifera (O.F. Müll.) J. Agardh 

Epicladia flustrae Reinke 

Epicladia perforans (Huber) R. Nielsen 

(§) Epicladia phillipsii (Batters) R. Nielsen 

Eugomontia sacculata Kornmann 

Gayralia oxysperma (Kütz.) K.L. Vinogr. ex Scagel et al. 

Gomontia polyrhiza (Lagerh.) Bornet & Flahault 

Monostroma grevilleï (Thur.) Wittr. 

Ochlochaete hystrix Thwaites in Harv. 

Ostreobium quekettii Bornet & Flahault 

Percursaria percursa (C. Agardh) Bory 

Phaeophila dendroïdes (P. Crouan & H. Crouan) Batters 

Prasiola stipitata Suhr in Jess. 

BANGIOPHYCEAE (RHODOPHYCEAE) 

# Acrochaetium reductum 

Epiphytic on Ceramium nodulosum and Polysiphonia 

brodiaei in the littoral zone at a sheltered and an exposed 

site. Monosporangia were observed in September. 

The species has been reported from the outer 

Oslofjord (Sundene 1953) and as Chantransia reducta 

Rosenv. from sector 8 (Levring 1937). 

§ # Aglaothamnion cf. pseudobyssoides 

Epiphyte on various algae at 20 m depth at a sheltered 

site. The plants were ecorticate, with main axes less than 

75 µm in diameter, and 55-74 µm at the base. Rhizoidal 

filaments were numerous in some plants, growing out from 

axial cells but not forming an adherent cortex (Fig. 2A). 

Sterile and tetrasporangial plants (Fig. 2B) were found but 

no developed female reproductive structures were found 

that could give a definitive identification of the species 

(L’Hardy-Halos & Rueness 1990). The tetrasporangia were 

ovoid 1.2-1.5 times as long as broad (37-48 × 27-41 µm). 

The highly similar A. tenuissimum (syn. A. byssoides (Arn. 

ex Harv.) Boudour. & Perret-Boudour., Furnari & al. 1998) 

found in the same area had main axes with a diameter of 

70-115 µm and none or a few rhizoidal filaments in part 

adherent to the axes. In this species young slightly lobed 

cystocarps were observed. A. pseudobyssoides has been 

reported from the south western part of the British Isles to 

Portugal (Maggs & Hommersand 1993). 

Audouinella efflorescens 

Syn.: Grania efflorescens (J. Agardh) Kylin 

Recorded in the sublittoral zone at a sheltered site. Filaments 

4-6.5 µm in diameter, cells 8-10 times as long. 

Tetrasporangia were present in February, karpo- 

Pringsheimiella scutata (Reinke) Marchew. 

Protomonostroma undulatum (Wittr.) K.L. Vinogr. 

(§) Pseudendoclonium dynamenae R. Nielsen 

Pseudendoclonium fucicola (Rosenv.) R. Nielsen 

Rhizoclonium implexum (Dillwyn) Kütz. 

Rhizoclonium riparium (Roth) Harv. 

Spongomorpha aeruginosa (L.) C. Hoek 

Sykidion droebakense Wille 

(§) Syncoryne reinkei R. Nielsen & P.M. Pedersen 

Tellamia contorta Batters 

Ulothrix speciosa (Carmich. ex Harv. in Hook.) Kütz. 

Ulva lactuca L. 

Ulvaria fusca (Postels & Rupr.) Rupr. 

§# Ulvella setchellii P.J.L. Dang. 

Uronema curvatum Printz 

# Urospora microscopica Levring 

Urospora penicilliformis (Roth) Aresch. 

sporangia in September. 

Records are rare (Sundene 1953; Sivertsen 1981), but 

the species is assumed to be distributed all along the 

Norwegian coast according to Rueness (1977). 

# Callocolax neglectus 

Found associated with Callophyllis laciniata at 25 m 

depth at an exposed and a sheltered site. Reproductive 

female plants observed in September. 

(§) # Ceramium botryocarpum 

Referred to this species were tufted plants with procumbent 

branches attached to the substratum by many rhizoids. At 

the nodes 6-7 periaxial cells were found. Tetrasporangia 

were observed in September. The species occurred in the 

littoral zone at exposed and sheltered sites. Known from 

the British Isles and France, wider distribution unknown 

(Maggs & Hommersand 1993). 

Ceramium cimbricum 

Occurred at 5-11 m depth at a sheltered site. Reported 

from two sectors in Norway, but expected to occur all 

along the Norwegian coast (Rueness & al. 1997c). Otherwise 

reported from Denmark to the Arctic (see 

Athanasiadis (1996) for references). 

# Ceramium diaphanum 

Syn.: C. tenuissimum (Roth) J. Agardh 

Epiphytic on Zostera marina and Asperococcus bullosus 

in the sublittoral zone at two sheltered sites. Tetrasporangia 

were observed in September. Reported under 

the synonym in older publications.


Fig. 2. Red algae. A, B. Aglaothamnion cf. pseudobyssoides. A. Main axes with rhizoids not forming a cortex (arrows). 

B. Tetrasporangium. C. Erythrodermis traillii, crustose stage with tetrasporangia. D. Schmitzia hiscockiana, crustose stage with 

tetrasporangia (arrow). E-F. Rhodochorton purpureum male plants with spermatangia. Scale bars A: 500µm. B-D, E: 50µm. 

F: 100µm. 

(§) Ceramium pallidum 

Plants referred to this species were found in the littoral 

zone at exposed sites. Cystocarps, spermatangia and 

tetrasporangia were observed in September. Common 

at the British Isles, appears to be widespread in the North 

Atlantic, known under various synonyms according to 

Maggs & Hommersand (1993). 

(§) Ceramium siliquosum 

A female specimen was found in the littoral at a sheltered 

site in September. Not earlier recorded in Norway 

under this name, but reports of C. diaphanum sensu 

Harvey may be this species, see Maggs & Hommersand 

(1993). 

# Colaconema nemalii 

Syn.: Chantransia thuretii (Born.) Kylin var. 

amphicarpa Rosenv. 

Epiphyte on various algae in the upper sublittoral zone 

at two sheltered sites. Plants referred to this species differ 

from C. savianum by the shorter monosporangia 

(< 20 µm ) placed in clusters. Monosporangia were observed 

in September. Published records are rare, and

46 Sarsia 84:39-53 – 1999 

have possibly been included in records of C. thuretii. 

Reported from sector 8 by Hygen & Jorde (1935). 

# Colaconema strictum 

Syn.: Acrochaetium strictum (Rosenv.) Hamel, 

Chantransia stricta Rosenv. 

Epiphyte on Polysiphonia elongata at 9 m depth at a 

sheltered site. Not included by Rueness & al. (1997c). 

Reported from sectors 1 and 8 by Hygen & Jorde (1935), 

Levring (1937), and Sundene (1953). 

# Compsothamnion thuyoides 

Epiphyte of Laminaria hyperborea and on hydroids 

from 12-28 m depth on a sheltered and a moderately 

exposed site. Tetrasporangia were observed in September. 

Only reported once in Scandinavia just south of 

the investigated area (Boye 1896). It was recently found 

in the northern part of the Kattegat, Denmark (RN, unpublished 

observations) and is, therefore, not included 

in Table 3 (see discussion below). 

Cryptopleura ramosa 

Epiphyte on haptera of Laminaria hyperborea. Occurred 

at 5 and 8 m at an exposed and a sheltered site. Reported 

from three sectors (1, 8, and 12) by Rueness & 

al. (1997c). 

Erythrodermis traillii 

The crustose tetrasporophyte was found at 12 m depth 

at an exposed site. Tetrasporangia were observed in February 

(Fig. 2C). The sporophyte has been reported once 

before in Norway (Marstein 1997) while gametophytes 

are reported from five sectors (Rueness & al. 1997c). 

Erythrotrichia reflexa 

Epiphyte on several filamentous algae and on the 

hydroid Dynamena pumila (L. 1758) in the littoral zone 

and down to 2 m at two sheltered sites. Gonidia were 

observed in September. This southern species has previously 

been reported from sectors 8, 13, and 16 

(Rueness & al. 1997c). 

(§) Gracilariopsis cf. longissima 

A single sterile specimen was found at 8 m depth at a 

sheltered site. The identification is preliminary; as for 

confirmation, observation of reproductive structures or 

DNA studies are necessary. Not earlier recorded under 

this name from Norway, but according to Steentoft & 

al. (1995) this species might have been included under 

Gracilaria confervoides (L.) Grev. or G. verrucosa 

(Huds.) Papenf. reported from several sectors along the 

Norwegian west coast by Levring (1937), Wennberg 

(1950) and Stokke (1957). Collections of G. verrucosa 

from Denmark and Sweden, located at the Botanical 

Museum (C) have recently been reinvestigated and identified 

with Gracilariopsis longissima (M. Steentoft, personal 

communication). Due to uncertainty about distribution, 

the species is not included in Table 3 (see discussion 

below). 

# Halurus flosculosus 

Found at 4-9 m depth at a sheltered site. This rare species 

has only been recorded a few times before on the 

south western coast of Norway as Griffithsia setacea 

(Ellis) C. Agardh (Hansteen 1892, Levring 1937). 

Lithothamnion norvegicum 

Found at 17-28 m depth at an exposed site. The specimens 

correspond very well with the syntype (L), 

designated by Woelkerling & Verheij (1995). 

Areschough (1875) originally described this entity as 

Lithothamnion calcareum (Pallas) Aresch. var. norvegicum 

Aresch. Recorded along the Norwegian west 

coast north and south of the investigated area (e.g. Foslie 

1895 (as L. coralloides f. norvegica (Aresch.) Foslie), 

Printz 1926, Levring 1937); not included by Rueness 

& al. (1997c). 

# Plagiospora gracilis 

Epiphyte on the stipe of Laminaria hyperborea in the 

sublittoral zone at a semiexposed site. Tetrasporangia 

were observed in February. 

# Pleonosporium borreri 

Sterile specimens were found at 25 m depth at two sheltered 

localities in September. The apical cells were about 

25 µm in diameter. This rare species has been reported 

only once before from Norway (Boye 1896) just south 

of the investigated area. 

Rhodochorton purpureum 

Syn.: Audouinella purpurea (Lightf.) Woelk. 

Common in the investigated area in the littoral zone 

and down to 28 m depth, often as an epiphyte on stipe 

and haptera of Laminaria hyperborea. Tetrasporangia 

were observed in February and at one site male specimens 

with terminal clusters of spermatangia (Fig. 2 E, 

F) were also observed, similar to those reported in culture 

by West (1969). Although the species has been reported 

from most parts of world (see Garbary (1987) 

for references) mature gametophytes have hardly been 

observed in nature (Dixon & Irvine 1977).

Rhodomela confervoides 

Includes specimens referred to f. virgata (Kjellm.) 

Rosenv. based on the presence of the characteristic 

adventitious tetrasporangial branches. These forms were 

collected at 12-28 m depth at sheltered sites in February. 

Previously this entity has been recorded as R. 

virgata Kjellm. from sectors north and south of the investigated 

area (Printz 1926, Levring 1937). 

Sahlingia subintegra 

Epiphyte on Sphacelaria caespitula and a hydroid at 

14-25 m depth at an exposed site. Previously reported 

from sectors 3 and 17 (Rueness & al. 1997c). 

# Schmitzia hiscockiana 

Crustose epiphytes on stipes of Laminaria hyperborea, 

recorded at 9-36 m depth at 5 semiexposed to exposed 

sites. The identification is mainly based on the observation 

of irregularly cruciately divided tetrasporangia 

(19-21 × 10-11 µm) in February (Fig. 2D), and the fact 

that cell fusions were absent in the basal layer. The 

distinctive large and small surface cells, usually found 

in S. hiscockiana were not present (Maggs & Guiry 

1985). This is the first record from the west coast of 

Norway; S. hiscockiana has been reported from the 

Oslofjord (Karlsson 1995). 

# Titanoderma laminariae 

Epiphytic on the haptera of Laminaria hyperborea 

found at 5 m depth at an exposed site. Tetrasporangia 

were observed in September. Previously recorded south 

of the investigated area (Kjøsterud 1997). 

FUCOPHYCEAE (PHAEOPHYCEAE) 

Chilionema ocellatum 

Epiphytic on the lamina of Laminaria saccharina at 

12 m depth in a sheltered site. The specimens were characterized 

by a uni- to di-stromatic basal layer and 

multiseriate plurilocular sporangia measuring 15-17 × 

55 µm. Reported from sectors 5, 6, and 12 (Rueness & 

al. 1997b). 

Desmarestia ligulata 

Well developed plants were observed at 5 m depth at 

an exposed site in September. Previously reported from 

sectors 7, 8, 12 (Rueness & al. 1997b). 

# Myriactula chordae 

Found at 3-5 m depth at sheltered sites in September 

epiphytic on Acrothrix gracilis, Chorda filum, and 

Spermatochnus paradoxus. The assimilatory filaments 

were fusiform with cells 12-25 µm in diameter and 1- 


1.5 times as long as broad. Hairs and unilocular 

sporangia were present. Earlier reported from sectors 

5, 6, and 8 (Rueness & al. 1997b). 

Myrionema foecundum 

Epiphyte on Palmaria palmata, in the littoral zone at a 

sheltered to moderately exposed site. The plants were 

in agreement with the description and drawings of 

Phycocelis foecunda Strömfelt (1888) and plurilocular 

sporangia were present in September. In referring the 

species to Myrionema, we follow the concept of 

Sauvageau (1897), who emphasized the upright filaments 

from each cell in the basal disc as a main character 

for the genus. 

CHLOROPHYCEAE 

§ # Acrochaete heteroclada 

Epi-endophyte of Chondrus crispus in the littoral zone 

at an exposed site and at 1 m depth at a sheltered site. 

The specimens were in agreement with observations by 

Correa & al. (1988) and the records were confirmed by 

culture studies. 

§ # Acrochaete inflata 

Syn.: Pseudodictyon inflatum Erce. 

Endophyte of Chylocladia verticillata at 3 m depth at a 

sheltered site. The relatively long celled narrow filaments 

(4-5 µm) within the cell wall of the host are very 

characteristic with branch angles close to 90° (Fig. 3A). 

A few tiny Acrochaete-hairs and sporangia with two 

different sized swarmers were observed in September 

(Fig. 3B, C). The species has been reported from several 

sites in the Mediterranean Sea (Gallardo & al. 

1993). Otherwise rare, but more widespread than published 

records indicate (RN unpublished observations). 

Acrochaete repens 

Includes specimens described by Printz (1926, p 240) 

as Entoderma reticulata (Syn.: Entocladia reticulata 

(Printz) Levring) as a consequence of culture studies 

on material collected close to the type site by Sivertsen 

(1981), (RN unpublished observations). In the present 

study a few colourless hair-cells were noted on the 

endophytic filaments between the epidermal cells in the 

lamina of Laminaria hyperborea and L. saccharina. 

Chlorocystis cohnii 

Syn.: Chlorochytrium cohnii Wright 

Found associated with “Schizonema-tubes” in a 

rockpool. Recorded from sectors 8, 12, and 13 (Printz 

1926, Levring 1937).

48 Sarsia 84:39-53 – 1999 

Fig. 3. Green algae. A-C: Acrochaete inflata in cell wall of Chylocladia verticillata. A: Branched filaments, intercalary sporangia 

with two different sized spores (arrows). B-C: Side view of filaments within the thick cell wall of Chylocladia verticillata. 

B: Sporangium with exit tube (arrow). C: Acrochaete-hair (arrow). D: Ulvella setchellii on Phyllophora crispa, bifurcate marginal 

cells (arrows). Scale bar A-D: 50µm. 

# Cladophora dalmatica 

Syn.: C. oblittorata J. Söderstr. 

Found at 2-4 m depth at two sheltered sites. 

Cladophora flexuosa 

Found in a rockpool and in the littoral zone of two sheltered 

sites. This species was included under Cladophora 

sericea s.l. by Rueness (1977) and Rueness & al. 

(1997a) but is now accepted as a separate species 

(Jónsson & al. 1989). Reported from sectors 1, 2, 8, 

and from sector 12 by Söderström (1963) who included 

specimens identified by Printz (1926) as C. crystallina 

(Roth) Kütz. 

# Cladophora pygmaea 

Found growing on pebbles at 2-14 m depth at sheltered 

and moderately exposed sites as well as and in a 

rockpool. Recorded from sector 2 by Sundene (1953). 

Epicladia perforans 

Syn.: Endoderma perforans Huber, Entocladia 

perforans (Huber) Levring 

Endophyte in epidermal cells of old leaves in Zostera 

marina at 2 m depth at a sheltered site in September. 

Sporangia were present. Reported from sectors 8 and 

12 (Kylin 1910; Printz 1926; Sundene 1953). 

(§) Epicladia phillipsii 

Endozoic within the theca of the bryozoa Alcyonidium 

hirsutum (Fleming, 1828) in the littoral zone and down 

to 12 m depth at exposed to sheltered sites. This is the 

first record from Norway, although it is likely that reports 

of Epicladia flustrae may also cover this entity, 

originally described as E. flustrae var. phillipsii Batters 

(see Nielsen 1984).

Eugomontia sacculata 

In empty mollusc shells at 8-28 m depth at several sites. 

Reported only once from Norway (sector 12) by 

Sivertsen (1981). 

Phaeophila dendroides 

Occurred as an epi- or endophyte associated with various 

algae and within bivalve shells in the shallow 

sublittoral zone at a sheltered site. Reported in 3 sectors 

(Rueness & al. 1997a). 

(§) Pseudendoclonium dynamenae 

Found endozoic within the theca of the hydroid 

Dynamena pumila in the littoral zone at exposed to sheltered 

sites. Sporangia were observed in September. Not 

earlier recorded in Norway under this name, but it is 

likely that reports of Epicladia flustrae associated with 

hydroids in the littoral zone represent this taxa (Kylin 

1910; Levring 1937). Separated as a distinct species by 

Nielsen (1984). 

Pseudendoclonium fucicola 

Syn.: Pseudopringsheimia fucicola (Rosenv.) Wille. 

Bas.: Ulvella fucicola Rosenv. 

Epiphyte on Fucus vesiculosus observed in the littoral 

zone at an exposed site. Reported from 3 sectors 

(Rueness & al. 1997a). Probably more widespread than 

published records indicate. 

Rhizoclonium implexum and R. riparium 

Unattached, unbranched, uniseriate filaments referable 

to Rhizoclonium were observed in several sites. There 

was a distinct difference between filaments 10-12 µm 

in diameter and others about twice as broad, even within 

individual collections, therefore we distinguished 

R. implexum with filaments 10-15 µm filaments from 

R. riparium with (16-) 20-35 (-40) µm filaments. In the 

genus Rhizoclonium some authors refer to R. riparium 

var. implexum (Dillw.) Rosenv. (e.g. Silva & al. 1996) 

or consider R. implexa, R. riparium conspecific with R. 

tortuosum (Dillw.) Kütz. (e.g. Burrows (1991) and 

Rueness & al. (1997a) who reported Norwegian records 

under the last name). Taxonomy and distribution still 

in need of critical revision. 

Sykidion droebakense 

Epiphyte on Rhizoclonium implexum and R. riparium 

in a rockpool and at 2-3 m depth at two sheltered sites. 

It was described by Wille (1901) from the inner 

Oslofjord and recovered by Sundene (1953) from the 

same area (sector 2). It was also found by Hygen & 

Jorde (1935) and Jorde (1966) in sector 8, and by Printz 


(1926) on the Halten archipelago, probably in the northern 

part of sector 12. 

(§) Syncoryne reinkei 

Epiphyte on Sphacelaria caespitula at 2 m depth at a 

sheltered site. Sporangia were present in September. The 

species has not previously been reported from Norway 

under this name, but could have been included under 

Pringsheimiella scutata from which it was separated 

by Nielsen & Pedersen (1977). 

Ulothrix speciosa 

Found at 2 m depth at a sheltered site. Referred to this 

species were smoothly surfaced filaments with a diameter 

of 15-17 µm, cell length 10-17 µm, and 2-3 

pyrenoids per cell. Older collections identified as U. 

flacca from sectors 2, 8, 11, and 17 were identified with 

Ulothrix speciosa by Lokhorst (1978). 

§ # Ulvella setchellii 

Epiphyte of Phyllophora crispa, forming nearly circular 

discs up to 350 µm in diameter. Characterized by 

the relatively long cells in the radiating filaments, and 

some bifurcate marginal cells (Fig. 3D). Common in 

the study area and recorded down to a depth of 25 m. 

# Urospora microscopica 

Epiphyte on various algae in rockpools and at 2-20 m 

depth at sheltered to moderately exposed sites. The 

plants were 3-10 cells long, the basal cell having a lobed 

holdfast. The filaments measured 12-20 µm in diameter 

and tapered toward the base. Terminal or subterminal 

sporangia were observed in September. The species has 

not been reported since the original description based 

on material from the Bergen area (Levring 1937). 

DISCUSSION 

The investigations (1994-97) have contributed much 

new information about the marine algal flora of the Norwegian 

west coast. Newly recorded species for Norway 

include Acrochaete heteroclada, 

A. inflata, 

Aglaothamnion cf. pseudobyssoides, and Ulvella setchellii. 

Also newly recorded are Ceramium botryocarpum, 

C. pallidum, C. siliquosum, however, future investigations 

may show them to be conspecific with algae 

earlier reported under other names. Epicladia 

phillipsii, Gracillariopsis longissima, Pseudendoclonium 

dynamenae, and Syncoryne reinkei have recently 

been established and may have been reported 

earlier under the name of the species with which they 

were combined. In addition to finding species new to 

Norway, the study has extended the known northern

50 Sarsia 84:39-53 – 1999 

boundary of distribution along the Norwegian coast for 

20 species. 

New records for the area, are 157 species of Bangiophyceae, 

Fucophyceae, and Chlorophyceae. A number 

of these were expected to occur due to known distribution 

north and south for the investigated area. Although 

there have been collections from different seasons and 

types of sites, many expected species (e.g. Porphyra 

linearis Greville) are still not found in the area, nor have 

we recovered Gloiosiphonia capillaris which was found 

by Boye about 100 years ago (The Botanical Museum, 

University of Bergen). More effort is needed both in 

the collection and identification of special groups such 

as the corallines, small endo- or epiphytic brown algae 

and specialized species (e.g. Fucus ceranoides L.), that 

only grow in sites with brackish water (Lein 1984). In 

summary, 33 Nostocophyceae, 121 Bangiophyceae, 64 

Fucophyceae, and 56 Chlorophyceae are known to occur 

in sector 10. 

Many species with a southern distribution reach their 

northern limit along the Norwegian west coast. This coincides 

with the general decrease in temperature (Sætre 

1973) which is the main gradient along the Norwegian 

west coast. The present study documents a number of 

such species, some of them have a peculiar distribution 

in Norway restricted to sectors 8-16 (Fig. 1). These are 

listed in Table 3, which also includes other species with 

a similar distribution. Included are also Lomentaria 

articulata although a small sterile plant has been reported 

from sector 17 (Kleen 1874). Species observed 

in the present study are marked with an asterisk while a 

parenthesis indicates uncertainly about distribution. 

Ceramium botryocarpum and Titanoderma laminariae 

Table 3. Algae reported only at the Norwegian coast from 

Bergen to Vega (sectors 8-16, Brattegard & Holthe 1997) and 

not elsewhere in Scandinavia. Species recorded in sector 10 

are indicated by *. Species that have an uncertain distribution 

in Scandinavia are given in parentheses. 

Aglaothamnion priceanum Maggs, Guiry & Rueness 

*(Ceramium botryocarpum J.W. Griff. ex Harv.) 

(Coccomyxa astericola Rosenvinge) 

Codium vermilara (Olivi) Delle Chiaje. 

Feldmannia simplex (P. Crouan & H. Crouan) Hamel 

Gelidium pusillum (Stackh.) Le Jolis 

* Halurus flosculosus (J. Ellis) Maggs & Hommersand 

Lomentaria articulata (Huds.) Lyngb. 

Myriactula stellulata (Harv.) Levring 

* Nitophyllum punctatum (Stackh.) Grev. 

* Pleonosporium borreri (Sm.) Nägeli 

*(Titanoderma laminariae 

(P. Crouan & H. Crouan) Y.M. Chamb.) 

*(Urospora microscopica Levring) 

may have been recorded under other names. Urospora 

microscopica and Coccomyxa astericola are small species, 

they are reported from sector 8, but may easily 

have been overlook in other sites. 

None of the species in Table 3 have been observed 

along the south-east coast of Norway or elsewhere in 

Scandinavia (Nielsen & al. 1995; Brattegard & Holthe 

1997; Tolstoy & Willén 1997). The distribution of these 

species appears to be correlated with the relatively warm 

Atlantic water reaching the coastline between Bergen 

(sector 8) and Vega (sector 16) in winter (Lee & Ramster 

1981), with the mean of the coldest month above 4.2 

°C (Midttun 1975). In general, the species in Table 3 

are distributed around the British Isles, especially on 

the south-west coast of England where the mean winter 

temperature is above 7 °C (Lee & Ramster 1981). In 

areas where the mean winter temperatures range from 

4-7 °C, such as in the southern part of the North Sea 

(Belgium, Netherlands), the east coast of England, Scotland, 

the Shetlands and the Faroe Islands (Lee & 

Ramster 1981) the species do occur but are rare (South 

& Tittley 1986; Fletcher 1987; Burrows 1991; Maggs 

& Hommersand 1993), and they have not been reported 

in Germany, Denmark, and Sweden where the mean 

winter temperature in the surface water falls below 3.5 

°C (South & Tittley 1986). 

The occurrence of rare southern species may not be 

constant even in a short term perspective. Higher temperature 

some years may favour the reproduction or 

growth of species that probably survive under 

suboptimal temperature conditions along the Norwegian 

coast. Apart from the lack of investigations of the 

area, normal winter temperatures and an extraordinarily 

warm summer in 1997 may have contributed to the 

observation of a large number of rare species, and to 

the extended distributional limits for southern species 

living at the fringe of their distribution range. A general 

increase in the temperature of the North Atlantic 

due to natural temperature fluctuations or human activity 

is expected to favour such species, but further experimental 

data are needed to evaluate the influence of 

temperature and other environmental factors (e.g. light 

conditions) on the distributional pattern of these species. 

ACKNOWLEDGEMENTS 

The authors are very grateful to colleagues who took part in 

the field work or assisted with identifications. Staffan Hjohlmann 

of Bergen gave valuable help in February and Mikaela 

Kruskopf of Bergen managed and participated in the diving 

operations in September. Juliet Brodie of Bath verified the 

species of Porphyra, Sigurdur Jónsson of Paris identified or

verified the species of Cladophora, Christine Maggs of Belfast 

identified the crustose Schmitzia hiscockiana, Jan Rueness 

of Oslo verified the species of Aglaothamnion and Ceramium, 

Poul Møller Pedersen of Copenhagen verified the species of 

Myrionema and Margaret Steentoft of Portsmouth identified 

Gracilariopsis longissima and Gracilaria gracilis. Susse 

Wegeberg of Copenhagen identified and contributed the 

Corallinales. The crew of R/V Hans Brattström: Tore Hegg- 

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Accepted 1 September 1998 – Printed 16 April 1999 

Editorial responsibility: Ulf Båmstedt

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