The algae of the Arctic Sea, a survey of the species, together with an exposition of the general characters and the development of the flora
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Vi, SEA eeeGes, . | eens Ag), ALP cba bat, Ltcetclates (ll) fseates De A capa will gitar? 1 = lac] 3 el > 3 a> o a Pe 2 5 ‘a Do Sb Eg eo © Oy a 5 5 mee 8 PS ne w ne nm Ss ot © Be 7 £ PE S a), Sects Uh leeccee jl peace 1 1 aes lite ewe wee NN Seewene 1 ati, Sie | acoSeees |! faateteeas ‘ih | Beane 1 1 1 DS oll) steeds Wilt beteoetl epebee |) -Seeceml 4 eteame:. IP) aeeess 1 seaeme-. |) mete Wy! Seema | oskeawne 1 48 KJELLMAN, THE ALG OF THE ARCTIC SEA. The history of the Flora of the Arctic Sea. It is seen from table 1 given above, that of the alge known from the Arctic Sea the following ones are not to be found south of it, as it has been defined here: Lithothamnion soriferum, Scytosiphon attenuatus, » alcicorne, Phloeospora pumila, » glaciale Dictyosiphon corymbosus, » intermedium, » hispidus, » flavescens, Sphacelaria arctica, » foecundum, Ectocarpus Lebelii, » compactum, Pylaiella nana, Lithophyllum arcticum, » varia, Polysiphonia Schibelerii, / Gleothamnion palmelloides, Delesseria rostrata, ’ Chetophora maritima, » corymbosa, » pellicula, Hemescharia polygyna, Ulva crassa, Halosaccion saccatum, Monostroma undulatum, Kallymenia septemtrionalis, » lubricum, » Pennyi, » cylindraceum, » rosacea, » saccodeum, Phyllophora interrupta, » angicava, Rhodochorton intermedium, » arcticum, » spinulosum, » leptodermum, » spetsbergense, » crispatum, Diploderma amplissimum, Rhizoclonium pachydermum, » miniatum, Chetomorpha Wormskioldii, Porphyra abyssicola, » septemtrionalis, Scaphospora arctica, Ulothrix Sphacelariz, Alaria membranacea, » discifera, » grandifolia, Characium marinum, » dolichorhachis, Codiolum Nordenskidldianum, » — elliptica, Chlorochytrium inclusum, » oblonga, Chlorangium marinum, Laminaria atrofulva, Rivularia microscopica, » nigripes, Gleocapsa spec. Lithoderma lignicola, Accordingly no less than 63 species, representing 22 families and 34 genera. A considerable number of these alge are, indeed, now described for the first time, or KONGL. SV. VET. AKADEMIENS HANDLINGAR. BAND. 20. n:0 5. 49 have only lately been distinguished as separate species, and it is thus certainly possible that some one or other of them, attention being now drawn to it, may prove to go southward of the limit established for the Arctic See. But on the other hand, the great majority consists of species so well marked by size and specific characters, that it seems rather improbable, if they did occur farther to the south, that they would have escaped observation on the coasts comparatively so accurately and long investigated of the northern Atlantic, where in such a case most of them would be expected to grow. About one third of them belong exclusively to those parts of the Arctic Sea which are not filled with ice: the Norwegian Polar Sea, the western Murman and the White Seas, namely: Lithothamnion soriferum, Monostroma undulatum, » alcicorne, » cylindraceum, » intermedium, » saccodeum, Polysiphonia Schibelerii, » angicava, Halosaccion saccatum, ° » arcticum, Diploderma amplissimum, » crispatum, Lithoderma lignicola, Ulothrix Sphacelarie, Ectocarpus Lebelii, Chetomorpha septemtrionalis, Pylaiella nana, Chlorangium marinum, Gleothamnion palmelloides, Gleocapsa spec. Cheetophora pellicula, Accordingly 21 species, representing 11 families and 14 genera. The other 42 species occur all of them in the Arctic Sea proper. Only 11 or 10 of these have been as yet met with also in the Norwegian Polar Sea, in the western Murman Sea, and in the White Sea. These are: Lithothamnion glaciale, Sphacelaria arctica, » flavescens, Pylaiella varia, Kallymenia septemtrionalis, Ulva crassa, Porphyra abyssicola, Monostroma lubricum, Alaria membranacea, Codiolum Nordenskidldianum. » grandifolia? Thus there remain no less than 81 or 82 species that are endemic in the Arctic Sea proper, namely: Lithothamnion foecundum, Rhodochorton spinulosum, » compactum, » intermedium, Lithophyllum arcticum, » spetsbergense, Delesseria rostrata, Diploderma miniatum, » corymbosa, Scaphospora arctica, Hemescharia polygyna, Alaria grandifolia? Kallymenia Pennyi, » dolichorhachis, » rosacea, » elliptica, Phyllophora interrupta, » oblonga, K. Vet. Akad. Handl. Bd 20. No 5. 7 50 KJELLMAN, THE ALG OF THE ARCTIC SEA. Laminaria atrofulva, Monostroma leptodermum, » nigripes, Rhizoclonium pachydermum, Scytosiphon attenuatus, Ulothrix discifera, Phloeospora pumila, Chetomorpha Wormskioldii, Dictyosiphon corymbosus, Characium marinum, » hispidus, Chlorochytrium inclusum, Chetophora maritima, Rivularia microscopica. These species belong to 15 different families and 22 different genera. This strong endemism points tothe purely arctic marine Flora being, contrary to the arctic phanerogamous Flora, no immigrated Flora, but one that possesses its centre of development in the Arctic Sea itself properly so called. Other circumstances lead cogently to the same conclusion, indicating at the same time that the present purely glacial marine Flora must have been formerly more widely spread towards the south than it is now. This results, I think, from a comparison of the Flora of the Arctic Sea with that of the northern Atlantic and the northern Pacific. The Arctic Sea, taken in a wide sense, possesses, as is shown by table 1, a con- siderable number of species in common with the Northern Atlantic. These species amount to 184 (185). By far the greatest part are met with on the Atlantic coast of Europe, only 11 are exclusively American. Amongst these, the following four are not known south of New-Foundland: Delesseria Montagnei, Fucus miclonensis, Laminaria Agardhii. Phyllaria lorea. Besides, I think it doubtful whether that plant from New-Foundland which has been called Laminaria caperata is in reality identical with the Laminaria Agardhi of the Arctic Sea. If this is not the case, Laminaria Ayardhit is to be reckoned amongst the endemic species of the Arctic Sea. The other 7 species: Ptilota pectinata, Antithamnion Pylaisei, Fucus edentatus, » evanescens. Laminaria longicruris, Agaruin Turneri, Phyllaria dermatodea either are not known south of Cape Cod or at least have their proper area of distri- bution north of this promontory *) and grow there most abundant and most richly developed. It is known already by the researches of Harvey, fully confirmed by later observations, that this promontory on the east coast of America forms the boundary between a more southern Flora and one expressly named arctic by American algologists °). 1) Cp. Fartow, New Engl. Alg. *) Fartow, |. c. p. 4. KONGL. SV. VET. AKADEMIENS HANDLINGAR. BAND. 20. N:0 5. 5] It cannot well be doubted that the real origin of the last-mentioned 7 species is to be placed in the Arctic Sea. Their occurrence on the coast of America south of the Arctic Sea is obviously to be explained by the Labrador current flowing along the coast from the north; it may have carried them down, and it makes moreover the external conditions similar to those under which these alge live in the Arctic Sea. Some of these species, Ptilota pectinata, Fucus evanescens, Laminaria longicruris, Agarum Turneri, and Phyllaria dermatodea, belong to those arctic species which are most commonly dispersed and occur in the greatest masses. They are all found in Baffin Bay, even at high latitudes. Delesseria Montagnei appears also to be a common species in Baffin Bay. At least it is to be found in quite considerable numbers in the collections from Greenland that I have had the opportunity of examining. Fucus edentatus and Anti- thamnion Pylaisei are also recorded from the same part of the Arctic Sea, but it is still unknown whether they are abundant or not. Thus there remain only two species that have not as yet been found with certainty in the Arctic Sea directly north of their reported American locality: Fucus miclonensis and Phyllaria lorea. The former species, as I understand it, is no properly arctic alga, the latter is on the contrary known from the most arctic parts of the Polar Sea. It is possible that Fucus miclonensis, both that which I have set down under that name from the Norwegian Polar Sea and that which J. G. Acarny reports from New-Foundland, Spitzbergen, and Greenland, is nothing else than one of the numerous forms in which Fucus evanescens presents itself, or else that Fucus miclonensis from New-Foundland is, in accordance with the opinion of J. G. AGarDH, identical with that form of Fucus, brought home from Greenland and Spitz- bergen, which I think ought to be regarded as a variety of the Fucus evanescens that is commonly dispersed in the Arctic Sea. With regard to Phylloria lorea, I can see no probable reason why it should be supposed to have had its centre of development at New-Foundland or to have come there from the south. On the contrary it may be assumed with very great certainty to have had its origin and centre of development in the Aretic Sea, just as the other species in question. With the exception of these species, the others possessed by the Polar Sea in common with the northern Atlantic are known either exclusively from the European coast of the Atlantic or both from here and from the north-eastern coast of America. Amongst these not a few are found in the purely arctic parts of the Polar Sea, reaching here very high latitudes, being widely distributed, occurring often in large numbers, in short, numbering among the most characteric alge of the Arctic Sea. On following their distribution southwards, some of them are found to disappear immediately south of the limit of the Polar Sea, others go only a few degrees of latitude south of the Polar Circle, others again, though common and luxuriant in the arctic waters, become southwards more and more rare, small and poor. These circumstances apparently in- dicate that those species have had their origin in the Arctic Sea and spread from here to the more northern parts of the Atlantic. The following alge may be quoted as good instances of such species: Halosaccion ramentaceum, known from all parts of the Arctic Sea, very common in certain regions, for instance in the eastern part of the Greenland Sea. It goes north- 52 KJELLMAN, THE ALG OF THE ARCTIC SBA. wards at least to the 80:th degree, and between the 79:th and 80:th degrees is still highly luxuriant. This is the case also with Polysiphonia arctica. In the European part of the Atlantic these two alge are not known south of Iceland; on the coast of America the former is one of the more prominent elements of the so called arctic Flora, occurring in large numbers at Eastport, and being only occasionally met with so far to the south as the coast of Massachusett '). Ralfsia deusta, though less common in the Arctic Sea, is yet found rather gene- rally in the most northerly part of the Murman Sea. In the Atlantic it has about the same southern limit as the preceding, being known from Iceland, but not south of Eastport (Maine) on the coast of America. Monostroma Blyttii in the Arctic Sea at Spitzbergen ascends to the 79:th degree of latitude; however, its maxima of frequency are on the north coast of Norway and on the south-western coast of Greenland. It is not known south of Iceland. On the north-western coast of America, it is found most abundant and luxuriant about East- port, descending southwards to about Boston ’). Rhodophyllis dichotoma, another characterical arctic alga, found on the coast of Spitzbergen about 79° N. Lat., is rather common in the north-eastern part of the Murman Sea, and common on the north coast of Norway. It is known from Feroe Isles and the neighbourhood of Bergen on the west coast of Norway, but not farther to the south on the European side. On the west coast of America its southern boundary is at Cape Ann. The following species: Odonthalia dentata. Rhodomela lycopodioides, Euthora cristata, Ptilota plumosa, » pectinata, Phloeospora tortilis, Cheetopteris plumosa, may undoubtedly be reckoned among the most characteristic species of the arctic Flora. In the European part of the Atlantic no one of them goes to the south of England, most of them are limited to the west coast of Scandinavia and the most northerly parts. of Great Britain. Those of them which are met with on the east coast of America have here a northerly distribution, belonging exclusively or chiefly to the arctic part of that region. Delesseria sinuosa,- Dichboria viridis, Desmarestia aculeata, Chetomorpha melagonium, all of them common species even in the most northerly parts of the Arctic Sea, are indeed to be found even south of England, but here they appear to be rare or even very rare. At least they are so at Cherbourg, according to the statements of Le Jozis a This is perhaps the case also with several other species, but it impossible at present ') Cp. J. G. Acarpu, Spec. Alg. II. p. 359 and Fartow, New Engel. Al *) Cp. Kssniman, Isl. Alg. p. 79 and Fartow, lL. ec. p- 5 and 42, S eM ee 3) Cp. Le Jouis, Liste Alg. Cherb. KONGL. SV. VET. AKADEMIENS HANDLINGAR. BAND. 20. N:o 5. 53 to arrive at any higher degree of certainty on this point, because the existing state- ments on the distribution of alge south of the Channel are scanty and fluctuating. Another reason which seems to speak for the Arctic Sea being an independent centre of development, is the fact that in the Arctic Sea and even in the parts of it situated to the north of the Atlantic there are found some species which are wan- ting in the Atlantic, but occur in the northern part of the Pacific. Such are: Delesseria Beerii, Petrocelis Middendorfii, Rhodymenia pertusa, Sarcophyllis arctica, Antithamnion boreale, Laminaria cuneifolia? » fissilis, » solidungula, Elachista lubrica. The first of these alge, as is shown by the table of the distribution of the spe- cies, is known from the Greenland Sea, and from the White and the Murman Seas. In the Greenland Sea it ascends to the north coast of Spitzbergen'), though it seems to be rare here; in the eastern Murman Sea it is more common and developed luxu- riantly ”), in the White Sea and the neighbouring part of the western Murman Sea it is one of the commonest alge’*). Without the limits of the Arctic Sea it is known with certainty only from the Ochotsh Sea. Another locality reported is Kamtshatka*). If we should assume that the centre of development of this species were the Pacific, it would become very difficult, not to say impossible, to explain its occurrence in the Arctic Sea north of the Atlantic. It cannot well be supposed to have migrated along the shores, nor to have been carried to its present place by currents. For the current goes from the west towards the east along the coast of Siberia, as has been proved by the researches of the Vega expedition. The alga could possibly have been brought to the American Arctic Sea by the Kurosivo-current, of which at least a feeble branch flows in that direction, but there is no current leading over to the Spitzbergen Sea from the American Sea. If on the contrary we suppose that it has originated in the Arctic Sea, there offers a probable and rather easy explanation, which I shall set forth below, to account for its occurrence in the Ochotsh Sea. It may certainly be objected that the species is known neither from the Siberian nor from the American Arctic Sea; but on the other hand it may be rejoined, that the American Arctic Sea and even the Siberian Sea are little known with regard to algology, and that therefore the species may very well exist in these seas. This is especially probable with regard to the Ame- rican Arctic Sea, because there is found in Baffin Bay a species very nearly related to and very slightly differentiated from D. Berti, namely D. corymbosa, which may pos- 1) Kistuman, Spetsb. Thall. 1, p. 12. 2) Cp. Kszuiman, Algenv. Murm. Meer. p. 13. 3) Goi, Algenfl. Weiss. Meer. p. 11. 4) G. Ruprecut, Alg. Ochot. p. 239 follow. 54 KJELLMAN, THE ALG OF THE ARCTIC SEA. sibly have issued from D. Berii or from some parent form common to both. Besides, there can be adduced no reason why the arctic alge might not be supposed to have been more widely distributed formerly in the arctic regions than they are now, which is assumed on very good reasons to be the case with the arctic phanerogams. What has been said now about Delesseria Berti, may be applied, though with one or two modifications, even to the other above-mentioned species, that are met with in the Arctic Sea and south of it only in the northern part of the Pacific. By means of the table exhibiting the geographical distribution of the arctic alge and the more detailed data set forth in the special part of this work, every one may easily convince himself of this fact. It is neither necessary nor suitable to enter here into details with regard to each species. But I will, however, call attention to one of them, Laminaria solidungula, one of the most peculiar and characteristic alge of the Arctic Sea. It is found in the Arctic Sea almost circumpolar: in the Greenland Sea, where it 1s com- monly distributed along the whole west and north coasts of Spitzbergen, and not seldom attains such a considerable size as to become one of the most magnificent alge of the ocean, in the Murman Sea, the Kara Sea, the Siberian Sea not far from the mouth of Behring Strait, and in Baffin Bay on the west coast of Greenland. It probably grows also in the American Arctic Sea. South of the Arctic Sea it is known only from one place, namely the Ochotsh Sea, from where Ruprecut reports a young »abnor- inal specimen of Laminaria saccharina with undivided scutiform root», which is in all probability a young Laminaria solidungula *). The present Flora in the nortiern part of the Pacific differs so essentially in composition from that of the Northern Atlantic, that is to say, it contains many species that are so sharply distinguished from those of the Atlantic, even belonging to quite different types, that in order to account in any way for this fact, one is necessarily obliged to assume that these two divisions of the ocean appertain to different areas of development, within which different forms have continued to be evolved during a very long time. However, on the other hand, it is a well-known fact that the northern Atlantic has no inconsiderable number of species in common with the northern Pacific. Though it is highly probable, as J. G. Acarpu has rightly remarked, that »a great part of the statements about alge occurring in widely distant seas is attributable to imperfect knowledge and wrong determinations caused thereby, and that the number of such species as are supposed to grow in widely distant seas will be diminished in proportion as the accuracy of the scientific determinations increases», and though this general judgment may be true even with regard to the reported number of species reputed common to the northern Atlantic and the northern Pacific, still there are un- doubtedly to be found in these widely separated sea-regions, as the same algologist expressly points out, several forms that can be proved to be identical ®). The present hydrographical conditions being so essentially unlike, it can hardly be assumed that these species should have been developed both in the northern Atlantic and in the 1) Cp. Ruprecut, Alg. Ochot., p. 351 and J. G. Acarpu, Lamin. p- 8 and Grénl. Lamin. och Fue. p. 11. A a. G. AcarpH, Spetsb. Alg. Progr. p. 1. Spetsb. Alg. Bidr., p- 10. Grénl. Lamin. och Fue. p. 8— : ite: KONGL. SV. VET. AKADEMIENS HANDLINGAR. BAND. 20. N:0 5. 55 northern Pacific, after these seas had received their present limits and physical nature ’). It seems as if J. G. Acarpa inclined to the opinion that these species common to both the regions in question have had their origin in either of them and have been brought from this region into the other by a current continued through all the Arctic Seas. AGARDH says: »Though it appears to be true in general with regard to the algw, that the distribution of a species is confined to the limits of the same sea-current, still the fact of many larger alge occurring both in the Atlantic and the Pacific may possibly indicate the existence of a current continued through the Arctic Seas, which might carry algee from New-Foundland and Spitzbergen to Kamtshatka and the most northern isles of Western America» ”), The hydrographical researches, carried on in the Arctic Seas by the Polar expeditions of later times, have not, however, demonstrated the exi- stence of such a continued current, having rather proved, on the contrary, that there is in the Arctic Sea a whole net-work of currents. Only by supposing a species to have been removed from one current into another, its occurrence both in the Pacific and the Atlantic could be explained from the influence of sea-currents, and such a combi- nation would be so complicated with regard to several species as to be hardly admissible. It might be supposed, indeed, that such a current existed during previous periods; but I do not know af any reasons for such a supposition. Without entering into this question in general, I will only remark, that in my opinion the occurrence of several alge in the northern Atlantic at the same time as in the Pacific may probably be ex- plained by the former distribution of water and land and the different physical con- ditions of the seas in former times as compared with their present state. The study of the present distribution of the alge of the Arctic Seas leads to such a conclusion. According to pretty reliable statements, the following arctic alge are to be found in the northern Atlantic as well as in the northern Pacific. Corallina officinalis? *Lithothamnion polymorphum, *Odonthalia dentata, *Rhodomela lycopodioides, Polysiphonia parasitica? » urceolata? » fastigiata? » atrorubescens? » nigrescens, Delesseria alata? i » sinuosa, *Hildbrandtia rosea, Peyssonnelia Dubyi, *Rhodophyllis dichotoma, *Euthora cristata, 1) Cp. ENGLER, Pflanzenw. p. IX—X. ") J. G. Acarpu, Spetsb. Alg. Bidr. p. 10. Plocamium coccineum, *Rhodymenia palinata, *Halosaccion ramentaceum, Dumontia filiformis, Callophyllis laciniata ? *Ahnfeltia plicata, Gigartina mamillosa? Chondrus crispus? *Ceramium rubrum, *Ptilota plumosa, oo pectinata, Callithamnion polyspermum, » arbuscula, Antithamnion floccosum? * D americanum, 56 KJELLMAN, *Rhodochorton Rothii, Porphyra laciniata, *Fucus vesiculosus, » ceranoides, » — spiralis? » evanescens, » miclonensis? *Alaria Pylaii? *Agarum Turneri, Phyllaria dermatodea, Laminaria saccharina? * D longicruris, » digitata? » stenophylla, *Chorda filum, *Ralfsia deusta, *Chordaria flagelliformis, THE ALGH OF THE ARCTIC SEA. *Scytosiphon lomentarius, *Desmarestia aculeata, *Dichloria viridis, *Phloeospora tortilis, *Dictyosiphon foeniculaceus, *Cheetopteris plumosa, *Ectocarpus confervoides, *Pylaiella litoralis, Enteromorpha clathrata, » intestinalis, © » compressa, Ulva lactuca, *Monostroma fuscum. *Spongomorpha arcta, Cladophora glaucescens? *Rhizoclonium ripariuin, *Chetomorpha melagonium, *Elachista fucicola, » tortuosa, *Lithoderma fatiscens, *Urospora penicilliformis, *Tlea fascia, Bryopsis plumosa. Of these 70 species, no less than 41, viz. those marked with an asterisk, accordingly 584 per cent, are at present known with certainty from the arctic parts of the Polar Sea, and amongst these 41 species there are several of the most commonly distributed and most characteristic forms of that region of the Polar Sea which is rich in ice. As moreover many of them, as has heen shown above, are chiefly distributed northwards at least in the Atlantic — there are no detailed statements to be had for the Pacific — we are justified in placing their origin in a glacial sea and in assuming that they have passed from there to the northern Atlantic and the northern part of the Pacific. The percentage of arctic forms amongst those species which are reported common to the Arctic Sea and the northern part of the Atlantic and of the Pacific, is in all probability larger than what is indicated by the figures mentioned. For there is some reason to suppose that a rather large number of species from the northern part of the Pacific, which have been considered identical with forms from the Atlantic, will prove on closer examination to be specifically distinct or wrongly determined. ‘This is probably the case with those marked with a sign of interrogation in the above list’). Their number amounts to 16. If these are deducted, the arctic forms would constitute about 75% of the whole number of species that the Arctic Sea has in common with the northern part of the Atlantic and the northern Pacific. Of the 13 species then remaining, Porphyr ina e morpha intestinalis auct., E. clathrata auct. and Ulva a ce pee ele 1) In support of this supposition, I refer to the notices on these speci i i I t 7 I pecies which are given by J. G. i Spec. Alg., Epicr. and his treatises on the arctic marine Flora, by Ruprecut in ile. Ciohot ne ee in New Engl. Alg, by Harvey in Ner. Am. and Alg. Vanc., by Postets and Ruprecut in lll Alg KONGL. SV. VET. AKADEMIENS HANDLINGAR. BAND 20. N:0 5. 57 with on the west coast of Greenland, are cosmopolitan, if they are understood in that extensive sense still usual with algologists, and on this account afford us no clue for deciding the question of their origin. Thus it has been shown that the arctic marine Flora is rich in endemic species; that several species which go far northwards in the Arctic Sea and are widely distri- buted there, are only slightly spread southwards in the Atlantic; that there are a number of species in the northern part of the Pacific which occur also in the Arctic Sea, even in those parts of it which lie north of the Atlantic, but are wanting in the Atlantic itself; and that of the species comparatively very numerous, which the arctic Flora possesses in common with the northern Atlantic as well as with the northern Pacific, a very large proportion consists of such species as are met with in the Arctic Sea at high latitudes, amongst them several of the most characteristic forms of the Arctic Sea. I think I am justified in drawing from these facts the conclusion that the Flora of the arctic part of the Polar Sea is an old Flora and that it has developed within the Arctic Sea. But this being so, it ought to be explained why several arctic forms are met with at present south of the limits of the Arctic Sea, both in the Atlantic and in the northern part of the Pacific. The occurrence of several arctic alge on the north-eastern coast of America is easily explained by their having been carried there by the cold Labrador current, which moreover makes the external conditions on the last-mentioned coast very similar to those in which the alge live in the Arctic Sea. But along the European coast of the Atlantic no current runs down from the Arctic Sea; on the contrary, a current flows upwards into the Arctic Sea from the Atlantic. Even if the occurrence of several arctic alow on this coast might be explained by their having wandered southwards along the coast, from the Spitzbergen Sea and the Mur- man Sea along the land of the Cisuralian Samoyedes a. s. 0., this explanation cannot be applied to the species met with at Iceland, and on the coasts of Great Britain and France. The currents between the northern Pacific and the Arctic Sea are favourable for carrying alge into the Arctic Sea, but not for transporting them from the Arctic Sea into the Pacific. However, the causes of the present distribution of the alge in the great divisions of the Ocean need not nor ought to be sought for in the conditions now existing on the earth, any more than the causes of the distribution of the land-plants. Just as the phanerogamous Flora of Scandinavia contains se- veral elements which are remnants from that period when the glacial formation extended farther southwards than in our days, those arctic alge occurring in the 1) Compare this species in the special part. K. Vet. Akad. Handl. Band. 20. N:o 5. 8 58 KJELLMAN, THE ALGH OF THE ARCTIC SEA. northern Atlantic and the northern Pacific may have stayed there from those times when northern Europe was surrounded with a sea filled with ice stretching down to the present northern coast of France. It is not at all improbable that the Flora of this sea may have been similar to that of the present Arctic Sea. When the glacial formation diminished in the south, southern plants immigrated in the sea as well as on the land, dislodging the main mass of the glacial ones. However, some of these, being able to hold out the struggle against the new-comers, maintained themselves in their original home and have done so ever since. Even though, as I have tried to show above, the present poverty in alge in the eastern part of the Kara Sea and in the Siberian Sea may depend essentially on the unfavourable nature of the bottom and the slight salinity of the water, still it is probable that during the oldest glacial period, when these seas extended farther southwards than now, the configuration of the coast and the condition of the water were more advantageous for alge and that the marine vegetation of these parts of the Arctic Sea was not then so poor’). The occurrence of several species, as De- lesseria Berti, Rhodophyllis dichotoma, Petrocelis Middendorffi, Ptilota pectinata a. o., in the Ochotsh Sea, the Murman Sea and the Spitzbergen Sea, and their absence in the intermediate region of the arctic waters, may be explained by the hypothesis that these species formerly grew also in the last-mentioned sea-region, but succumbed afterwards, because the shore was moved northwards, partly through the formation of deltas, partly through a general rising of the land elevating the sea-bottom, which was formed of sand-banks, and because large rivers began to pour ever increasing masses of fresh water into the sea. If Halosaccion saccatum, reported from the White Sea, is really to be found there, it should probably be numbered among those species which have formerly been more widely spread along the coast of Siberia; for, even though it be not identical with any of the species of the Pacific, H. fucicola, H. hydrophora, or H. jirmum, these are nevertheless its nearest relations. The change suffered by the Flora in the northern Atlantic and the northern part of the Pacific, in proportion as the glacial formation retreated, took place even in the part of the Arctic Sea surrounding the coast of Norway. By the immigration of southern forms, glacial species were su- perseded or deprived of their predominating influence. The elements of the Flora were considerably increased, and its character was no longer marked by glacial forms, but by such as belong to the Atlantic. However it is not only into this part of the Polar Sea, most closely allied hydrographically to the present northern Atlantic, that southern species have probably immigrated in later times, or are now in course of immigration; but this is the case also with the Arctic Sea proper. Such a transplantation of alge into certain parts of the Arctic Sea is very much favoured by currents flowing in that direction. Thus, for instance, alge may be easily transported into the eastern part of the Greenland Sea to the coast of Spitzbergen by the Gulf Stream, and into the eastern part of the Murman Sea to the west coast of Novaya Zemlya partly by means of this stream, partly along the almost continuous coast from the Norwegian Polar Sea and the western Murman Sea. Between Spitzbergen and Norway at different latitudes I ') NorDENSKIOLD, Proven, p. 70—71. KONGL. SV. VET. AKADEMIENS HANDLINGAR. BAND 20. N:0 5. 59 have seen Ozothallia nodosa and Fucus vesiculosus drifting, and on the south coast of Spitzbergen I have found Ozothallia nodosa washed ashore, overgrown with Polysiphonia fastigiata. I have not myself found any of these alge growing in any of the numerous places on the coast of Spitzbergen, that I have had the opportunity of examining. They are however stated by others to occur there. If this be really true, they may be assumed to have been transferred there in later times by the Gulf Stream. Whether any species have made use of the convenient route along the coast to Waygats and Novaya Zemlya cannot indeed be decided with certainty. It is possible, however, that Cladophora ru- pestris, found at the south-western extremity of Waygats, and Spongomorpha lanosa, met with at southern Novaya Zemlya, may have done so. It is possible also that a transfer of alge has taken place formerly and is perhaps going on even at the present time by means of those vessels which depart yearly in great numbers from northern Norway for Spitzbergen and Novaya Zemlya and stay during the summer in the waters sur- rounding these islands, as well as by those which start regularly since a long time for Baffin Bay from more southerly regions. I have evidence of a transportation of this kind taking really place, but I do not know whether the alge thus brought over into the Arctic Sea have been able or are able to maintain themselves there. One of the vessels employed in bringing over the Swedish expedition of 1872 to Spitzbergen, some time after the arrival of the expedition, was found to be richly clothed with small-sized Enteromorpha compressa, at and below the water-line. This plant had probably been engendered by spores, which had attached themselves to the vessel in more southern regions on the coast of Sweden or Norway and developed afterwards during the course of the voyage. It is possible also that water-fowls, especially such as inhabit lagoons on the coast, may carry with them some alga or other from the south. Perhaps Rhi- zoclonium rigidum, which grows abundantly in the lagoons at Advent Bay in Spitzbergen, has come to the high North in this manner. The marine Flora on the coast of Greenland includes a pretty considerable number of species whose origin lies in a southerly direction, in the Atlantic. The number is so considerable and the habitat of the algee is such that they cannot be supposed to have been transported into those regions by the agency of either men or animals. Nor can these species have come into the arctic waters about Greenland by means of seacur- rents directly from the south, i. e. from the east coast of America, as the current here goes from the north southwards, and, moreover, several species reported from Greenland, are wanting on the American coast. This is the case with Hydrolapathum sanguineum, Pelvetia canaliculata, Nitophyllum punctatum, Furcellaria fastigiata, Callophyllis laciniata, Asperococcus bullosus, Stupocaulon scoparium, Enteromorpha tubulosa. I do not certainly fee] quite sure that all these species occur really on the coast of Greenland. But as I have myself seen specimens of some species, stated to have been collected at Greenland, and as experienced algologists allege having seen specimens of the other species from the same regions, I could not but quote them for the present among the Greenland alge. I cannot explain their occurrence at Greenland otherwise than by the hypothesis that they have arrived there from the east by Iceland, where at least some of them have been 60 KJELLMAN, THE ALG OF THE ARCTIC SEA. hitherto met with. To be sure, there is no current leading directly from Iceland to Greenland, but by that current which passes along the coast of Iceland in a north- westerly direction, they may possibly have been carried into the region of the great Polar current or Greenland current and then brought on by means of that to the coasts of Greenland. But if this can have happened with these species, it can, of course, have happened also with other southern forms occurring on the coast of Greenland. The immigration to Greenland would thus have taken place not from the south, but from the east, from the European part of the Atlantic by way of Iceland. A closer investigation of the marine Flora of Iceland would surely give several important results on this point. Even from the northern part of the Pacific alge have surely immigrated at later times. However, I can hardly cite more than one reliable instance of it. No species is known with certainty from the Siberian Sea, that must be supposed to have immi- grated there from the Pacific, if we do not assume, as certain or probable, that those alge, Ptilota asplenoides and a species of Laminaria nearly related to L. longipes, which are said to be found in the Siberian Sea at the mouth of the Lena, do really grow in the place from where they have been reported '). If they belong to the Siberian Sea, which I think rather doubtful, we cannot but suppose that they have come there from the Pacific, where at least Ptilota asplenoides is widely distributed and grows in great abundancy at several places. The direction of the currents between these two seas does not favour the importation of alge into the Siberian Sea; it is more favourable for an immigration into the western part of the American Arctic Sea. However, this sea is as yet very little known with regard to its algology. Nevertheless, among the species reported from there, there is one that may be assumed, on good grounds, to have immigrated from the south, namely Rhodomela larix, which is extensively spread in the Pacific and common even in the northern part of the Behring Sea. Whoever visits the coasts of Spitzbergen, cannot but notice the objects of different kinds washed ashore, a great part of which have apparently been carried there from the south, from the coast of Norway. At some parts of the coast they are more rare, at others they are to be found in very large masses. NorpENSKIOLD relates, for instance, that at one place on the coust of Spitzbergen pieces of pumice were found in so great quantities that a small sack could be filled with them. Many of these objects, as floats of tree, glass, and cork a. o., are of such a nature that algw, at least smaller ones, can germinate on them, and in the earlies stages of their development be brought by means of them from the coasts of Norway to the North *). Larger alge may be trans- ferred floating on the water by the Gulf Stream. Thus it seems as if the marine Flora of Spitzbergen might be expected to comprise a rather considerable number of southern alge, immigrated from the north-west and north coasts of Norway at later times. This however is not the case. By far most elements of the Spitzbergen Flora have at pre- sent such a geographical distribution that they must be considered to have their proper home in the Arctic Sea. I have thought fit to call attention to these facts, because 1) Cp. J. G. Acarpu, Grénl. Lamin. och Fuce., p. 7. 