WAL A 9088 00903 3242 —- Il 173C68 1912 SSP AWE ALA OA eo Ae ts i 3 rae ae ee pawsol as ‘PROCEEDINGS OF THE. RovaL TRISH. ACADEMY | —Vowme XXXI CLARE ISLAND ) SURVEY Pare 15 DUBLIN: HODGES, FIGGIS, -& CO. Lap | LONDON: WILLIAMS & NORGATE. | 1D an “gre: Price Five Shillings _ [For scheme of publication of this series see inside of wrapper] : and the adjoining mainland in the West of Ireland was carried out bye one hundred specialists during the years 1909, 1910, ‘and OTE } undertaken in order to furnish a study of a Born area of the west! ‘coast baie problems in Europe; and also as a See towards the 2 stud e island faunas and floras—a subject of wide and absorbing interest. The report when finished will provide the most complete survey of the. Fauna and Flora of a definite area which has ever been made. In sada systematic zoology and botany, special attention has been given to questions of geographical distribution, dispersal, and ecology, and to the influence’ ‘of human operations upon the Fauna and Flora. The complete report is estimated to run to about 1600 pages with 100 plates, and will cost 60/ or more, The whole series may be subscribed for at the reduced price of 50/- paid in advance, on receipt of which the reports issued up to the present i will be forwarded, and future parts sent as soon as puree ‘Bach Lek ih will also be sold separately. the British Isles, and at least 1,000 species new to Tetends. The list printed on pages 3 and 4 of this wrapper shows the scope 0 work, the present state of publication, and the prices of the ne published so far. an publishers or to Tue SECRETARY -Rovan aa) = UL aT 7e2r7/e Clare Island Survey. i> Ceo GO) 12 AT aN ie MARINE ALGAE. By A. D. COTTON. Prates I.-XI. Read June 24 and Novrwmer 11. Published Novewner 27, 1912. CONTENTS. PAGE PAGE PREFACR, 1} ie The Salt-marsh Formation, . 7 INTRODUCTION, 5 2 y. The Vegetation of River- 1. Definition of the nccak 2 mouths, . $3 2. Previous work in the Area, 2 | . The Vegetation of Brackish 3. Field-work and Assistance, : 4 Bays, : : : - 88 I. Tue Externat ConpiTIONs oF THE )III. Tur Ateat Frora, . ° o 7 AD AREA, 5 ‘ : 6 1. The Systematic List, ; . 90 1. Climatic Gondificns A 6 2. Notes on the List, . : , 102 2. Physical Nature of the Coast 3. Economic Species, . . 161 and Conditions of Exposure, . 9 4, The Flora of The Bills, . . 165 II. Tue Atcar VEGETATION, . : - 12) Ty. Tue Cuaracter and Composition 1. Historical Résumé, . : =. Le or THE Fiona, : 156 2, Areas and Units of Vegetation,. 14 he Notes on Novelties and Additions, 156 3. The Vegetation, . 17 2. The Character of the Flora, . 160 i. The Rocky-shore Formation, 17 | 3. Comparison with other Areas, . 166 (A) Exposed Coasts, . 19 4. Origin of the Flora,. . . 168 (B) Sheltered Coasts, . 450) yy, Bysniograrny, Bee a ice hoe Wil ii. The Sand and ae mud Formation, 3 61 PREFACE THE natural history Survey of Clare Island and. the adjoining mainland provided an opportunity of investigating one of the most interesting algological regions in the British Isles. Since the first decade of the last century, when Miss Hutchins collected in Bantry Bay, and more especially since the days when Harvey published his famous “ Phycologia l’ritannica,” the west of Ireland has always possessed a great attraction for the marine botanist. But although various collections and many gatherings of new and rare species have been dealt with, no systematic investigation of that area has been undertaken, nor has any list or general account of the algal flora been published. The selection of Clare Island as a centre fora detailed survey was therefore particularly satisfactory from an algological standpoint. Results for the systematist and student of plant-distribution were certain, and the B,1.A, PROC., VOL. XXXI, A 15 NG 152 Proceedings of the Royal Irish Academy. variety of ground in the neighbourhood provided ample scope for the modern study of plant-communities. The position of the island—off the coast of Co. Mayo, and in the Gulf Stream drift—ensured the presence of that southern element which gives the Irish flora its peculiar interest. The following report shows that the hopes raised have been amply fulfilled. ‘I'he flora proved quite as rich as was expected ; and practically all the species for which the west of Ireland is noted occur within the survey- area. Valuable systematic results have been obtained; and the variety of algal associations present rendered the study of the marine vegetation one of the most comprehensive that has yet been carried out. Important phytogeographic data also, necessitating the modifying of previous views, were secured. INTRODUCTION. 1.—DEFINITION OF THE AREA. Owing to the comprehensive nature of a natural history survey, the limits of the area investigated varied in different cases, since in some it was advisable to include a much larger area than in others. . With regard to the marine algae, it was thought best to limit the investigation to Clare Island and the neighbourhood of Clew Bay. By concentrating attention on a small district a more accurate knowledge of the true nature of the vegetation can be obtained, and the variety of ground presented in the area mentioned proved quite as much as was possible to work with a moderate degree of thoroughness. The Survey-area for the algae, therefore, consisted of Clare Island, the shores and waters of Clew Bay, and an extension to the north to include Bellacragher Bay and Achill Sound. S At the same time it appeared advisable not to overlook the interesting records from Roundstone, a locality only twenty-five miles south of Clew Bay. This district was carefully worked some seventy years ago by W. McCalla, and he collected several species which were not found in the Clare Island district. A brief visit was paid to Roundstone; and all records from that locality are included in the report. 2.—PREVIOUS WorK IN THE ARFA. According to the scheme of geographical areas devised by: Adams (see Adams, ’11, in Bibliography) the province of Connaught is divided into three sub-divisions. Co. Mayo (which includes the survey-area proper) forms sub- division C 2, and, according to an annotated copy of Adams’ Synopsis (Adams, Clare Island Survey—Marine Algae. 15 3 ‘08), kindly sent me by the author, possesses thirty records of algae. Co. Galway, on the other hand, which forms sub-division C1, has a much longer list, owing to the collections made at Roundstone Bay. For the third sub- division, C3 (ie., counties Sligo and Leitrim), there are practically no records. With regard to sub-division C 2, Wm. Thompson, the zoologist, collected a few plants in Clew Bay about 1840; and Harvey quotes one record— Asperococcus bullosus—in “ Phycologia Britannica.” In a report of a natural history excursion to Achill at Easter, 1898, H. Hanna deals with the marine algae (Hanna, 98). He records twenty species; and this constitutes the only account of the sea-weeds of the area. Adams, in his list, adds nine names to the flora of the sub-division, culled from various sources. A single species has been recorded from Clare Island, namely Codium tomentosum (vide H. H. Dixon, Ann. Bot., xi, p. 590). The Roundstone records date from 1808. Fucus Mackaw was described from that locality by Dawson Turner (“ Historia Fucorum,” Pl. 52); and it still remains one of the few stations for the plant in Ireland. In addition to the discovery of this species, Mackay noted the presence of &hodochorton floridulum, which Harvey mentions in the chapter on algae in Mackay’s “Flora Hibernica.” McCalla’s records follow. He thoroughly explored the shore, and supplied Harvey, as the pages of “ Phycologia Britannica” testify, with much valuable material both from Roundstone Bay and Birturbuy Bay. McCalla was also one of the pioneers of dredging amongst algologists, and by this method he obtained many interesting species. He commenced to issue a set of “ Algae Hibernicae,” but, owing to his untimely death in 1849, only two volumes were published. Harvey visited Roundstone at least once, and in later times, Johnson (93) and H. Hanna. Fosle also, the well-known specialist on calcareous algae, came over from Norway on purpose to investigate the Lithothamnia which had been described from Roundstone by McCalla and Harvey. An account of his trip was published in the “Irish Naturalist ” (99, p. 175). Foslie was keenly interested in the Lithothamnia of the British Isles, and it was proposed to send him for identification all the Clew Bay material. His premature death in 1910 came as a great shock. Three days were spent by me at Roundstone in September, 1911, and, with the exception of Codium amphibium, all the species for which that locality is famous were seen in their natural habitat. The total number of species listed by J. Adams for the sub-division C 1 is 162. Outside the province of Connaught, Valencia Harbour (in the extreme south-west) has been specially investigated by biologists, and Weiss, who dealt with the algae [’00], gives a list of eighty-six species collected by him during August and September, 1896, with notes on the same. A 2 Qe 15 4 Proceedings of the Royal Irish Academy. 3. FIELD-WoRK AND ASSISTANCE. My visits to the Survey-area extended over three years. In each year two visits, varying from a fortnight to three weeks, were paid to the island and mainland, whilst during the last year, an additional winter visit was accomplished. Owing to the uncertainty of communication, no attempt was made during this last trip to cross to the island, but since the conditions of the exposed mainland are similar to those of Clare Island, a fair idea of the winter aspect of the vegetation of the whole area was obtained. In_ addition, Mr. J. Adains, of the Royal College of Science, Dublin, visited the district in August, 1909, investigating the island and dredging in Clew Pay. The dates of the visits were as follows :— 1909. April 8-20; July 12-27. [August 20-27, J. Adams. ] 1910. June 16-July 1; September 50-October 15. 1911. February 15-21; April 28-May 15; August 17-September 4. About one-half the time of each trip was spent on the island, and the remainder either on the mainland or in dredging. Spring-tides were usually selected for the former locality, when exposed or semi-exposed shores were examined at low-water. As is well known, spring-tides are more useful in sheltered regions than in exposed, and a greater number of rare and sub-littoral species are doubtless uncovered during springs in Clew Bay than on Clare Island. But since these can readily be obtained by dredging, there was no object in reserving low-tides for mainland work. Dredging was carried out in February, May, July, and August. Tor the more open waters I had the advantage of the ss. “ Helga,” of the Fisheries Branch of the Department of Agriculture and Technical Instruction for Ireland. The outer part of Clew Bay was explored, and a number of hauls were taken further out, ie., to the south and west of Clare Island, and near The Bills, in depths ranging from 10 to 20 fathoms. A large amount of material was collected, and considerable light obtained as to the nature of the benthos, but for detailed study much longer time than was at my disposal was necessary. J. Adams also dredged from the “ Helga” in 1909, and two interesting algae were found only on that occasion. For the inner part of Clew Bay a small sailing-boat was used: and good work was accomplished by means of a hand-dredge in 1-6 fathoms. The “coral-banks” were investigated, and in quiet weather very much may be learned as to the nature of the flora. The February dredgings were useful in obtaining fruiting specimens of encrusting algae. In May the mass of vegetation is so great that it is impossible to do more than sample various localities. As the hauls ot July, 1909, had provided several rare and interesting Clare Island Survey—Marine Algae. 15 5 plants, a few days were set apart in August, 1911, to search for note- worthy absentees. Rough weather, however, set in, and dredging from a small boat was out of the question. With regard to the nature of the field-work, the visits of the first year were mainly devoted to the listing of species from all parts of the area, and to working out critical forms. The same study was continued in 1910, and the analysis of algal communities was also taken in hand. The third year was principally devoted to the search for species which were to be expected but had not been found, and to the completion of the work on algal communities. On each visit, moreover, a large amount of materia! was collected for the Royal Botanic Gardens, Kew, as some extra leave had been granted for that purpose. The time devoted to ecology necessarily robbed the systematic list of a certain number of names. Searching for rare or microscopic species invariably yields results in the way of new or little-known plants, but the records of minute species, such as Cyanophyceae, are not yet of much value for comparative purposes. Moreover, the amount of time and labour spent in working out microscopic forms is often very great, and out of proportion to the results obtained. Unfortunately there was no specialist for the Cyanophyceae (which are abundant in brackish parts of Clew Bay) who could be appealed to for aid. The list, therefore, can hardly be considered as exhaustive. With regard to the littoral flora, the island-list must be fairly complete; and on the main- land the additions during the last trip were chietly microscopic forms. — But in the sub-littoral region there are probably many species yet to be discovered, as the vegetation is very local and less easy of exploration. The fact that such conspicuous weeds as Dictyopteris, Acrothrix, Dasya corymbosa, and several others were only seen once, confirms this view. Assistance.—Dr. P. Kuckuck joined the collecting party during June, 1910, and advantage was taken of his critical knowledge of the Phoeophyceae to work out Ectocarpi, Chordarieae, &c., and to make comparisons between the flora of the North Sea and Baltic, and that of Clare Island. In April, 1911, Dr. F. Borgesen paid a visit to the Survey-area. Naturally the vegetation of Clare Island had been continually compared with the detailed account given by him for that of the Faerées, and several striking contrasts had been noted. It was, therefore, of the greatest value to be able to examine the algal associations with Dr. Borgesen, and to discuss the problems that had arisen. His help also with regard to the Iceland flora was particularly useful. In order to make the floristic section of the work as accurate as possible, 15 6 Proceedings of the Royal Trish Academy. which for comparative purposes is of the very greatest importance, I have availed myself freely of the help of specialists. Dr. Kuckuck kindly worked out the Ralfsiae, and checked a number of other members of the Phoeophyceae. The weight of his authority in doubtful cases was most valuable. I am also much indebted to Dr. M. Lemoine. Since the death of Foslie, the identification of encrusting Lithothamnia was uncertain work, owing to the confusion in the literature, and the scarcity of reliable material in British herbaria. Madame Lemoine willingly undertook to examine the Trish material, and, by her method of anatomical investigation, to work out and compare critical species with authentic specimens. Though no important additions to the list have resulted, it is highly satisfactory to have trust- worthy identifications. An interesting point with regard to geographical distribution has, moreover, been cleared up by her. Dr. K. Rosenvinge kindly determined the Chantransiae, whilst Prof. N. Wille, Prof. C. Sauvageau, and Dr. H. H. Peterson examined the species of Ulothrix, Spacelaria, and Ceramium respectively. Prof. G. 8. West’s opinion on the brackish and freshwater species, and Mr. J. Adams’ aid in connexion with the distribution of Irish plants were most helpful. Mr. E. M. Holmes has taken the keenest interest in the Clare Island flora, and throughout the work of identification has always most liberally bestowed the benefit of his valuable and unique experience. ‘I’o all these botanists I offer my sincerest thanks. Special acknowledgments are due to Mr. R. Lloyd Praeger, the organizer and secretary of the Survey, to whom I am deeply indebted for advice on matters botanical and geological, as well as for encouragement and assistance in a number of ways. I.—THE EXTERNAL CONDITIONS*‘OF THE AREA. 1.—CLIMATIC CONDITIONS. 1, Atmospheric Temperature und Moisture.—The insular climate of Ireland, and the influence of the Gulf Stream drift, combine to render the Clare Island district particularly mild and damp. In winter rough stormy weather prevails, together with a large amount of rain, but frost is rare. In summer the weather is changeable, rain and gales from the south-west are frequent, and the average temperature relatively low. Though occasional warm days occur, spells of het dry weather are most unusual; and this renders the intertidal region more favourable for vegetation than that of the south coast of England. A glance at the isotherms of Europe shows that the mean January temperature of 45° F. (=6:11° C.) corresponds with that of the south of France and the north of the Adriatic; whilst the July isotherm of 58° F. Clare Island Survey—Marine Alyac. 15 7 (= 1444°C.) runs up the coast of Norway, across the north of Sweden to northern Russia and Siberia. Clare Island is situated on the same latitude as Heligoland and Kiel, but, as will be seen later, the flora is of a very much more southern type. The atmospheric moisture is of importance in connexion with the littoral vegetation. But the problem of desiccation during low water is a very complicated one, the latitude, together with the amount of wind and sunshine, having to be taken into consideration. The following figures (“Irish Coast Pilot,” ed. 6, 1911) give the mean humidity for the Blacksod Point Observatory for the last thirty-five years. It will be noted that the highest figures are those of the winter months, whereas in the Faerées the reverse is the case, for which the absence in Ireland of summer fogs is probably responsible :— | Jan. Febr. | Mar. | Apr. | May. | June. | July. | Aug. | Sept. | Oct, | Nov. Dee. | PESSmricSi le SG: o|seSban|i SI oul Nore.—Observations made at 8 a.m. 100 = Saturation point. | | | sl | 84 S5)))| 784 SL | ee | oe 2. Temperature and Salinity of the Sea.—Important as are atmospheric conditions, the temperature of the water is even more intimately connected with the algal vegetation. The direct influence of the former is confined to the intertidal region, and is thus intermittent; but the effect of the latter is much more general, being felt both by the littoral and sub-littoral flora. A complete series of observations on sea-temperatures at different depths is not available for Clare Island ; but the Meteorological Office has favoured me with details of the mean monthly surface-temperatures for Blacksod Bay and Cleggan during the year 1911; and Mr. G. P. Farran has kindly supplied me with two sets of. temperatures and salinities for various depths in Clew Bay and off Clare Island. From these it is evident that the surface temperature does not differ much from that of 10-20 fathoms, so that the monthly table given below affords a good general guide. The Blacksod and Cleggan observatories are about twenty miles north and south respectively of Clare Island. It will be noted that many of the Blacksod figures are slightly higher than those of Cleggan, for which local topography doubtless accounts :— MEAN SURFACE TEMPERATURE, BLACKSOD AND CLEGGAN, 1911. (C°). Teer hates eae ean | | Sar: | Febr. Mar. | Apr. | May. | June. | July. | Aug. | Sept. | Oct. | Nov. | Dee. | ——_—_-—_} |————__—— — — A | | Blacksod,| 7-2 | 6-7 | 6-7 | | 17 | 13-3 | 161 | 161) 13:3 6 | | 8 | } Cleggan, 72 | 67 67 | 83 10°6 | 7:2 67 10°6 | 7-2 5-6 11-1 | 12:2] 13-3 15:0 | 13:9 | 15 8 Proceedings of the Royal Irish Academy. CLARE ISLAND, May, 1909. i Surface. | 1-10 fathoms. 11-20 fathoms. |, 21-30 fathoms. 4 — | Station Number. Date. | | | | I Temp; | Sal. °/, | Temp.°C. | Sal. °/, || Temp.°C. | Sal. °/, || Temp. °C. | Sal. °/, ———- + -- —_—-- | sl Fide 4 —— = Clare Island W. 67, 21.5.709 | 11°15 34-61 SS — | (10-69 34-6401] | (20 fath | W. 69, | 22.5. 09 |) 11°35 | 34-60 = — |.. ==) °) = ipa Sie | | ( 24 fath . W. 70, . | 11°30 | 34-61 = de pane 34:67 || = | | | (18 fath. | \ } | 5 W. 72, ” 113 | 84°69 | (11-28 | 84-72 | — = he = | ( 9 fath. i | I ; W. 75, | 24.5. 709 || 11-0 | 34:74 = = ee | 34°72 | (10°55 | 84-72 | 12 fath. | {35 fath. ; W. 81, | 25.5. '’09|| 11-05 | 34-70 | eee | 1h 10:87 | | | ( 26 fath en MaeAWWE.82; - Ici aarogle e || ee eae a = | | | CLARE ISLAND, AuGustT, 1911. er oe Laare ester eet Rn eo nie, | i Clare Island W. 197, 18.8. ’11 || 17-40 | Bei fp = | = ae 340720 = | \| | | 18 fath, | \ “4 = - | 16:96 | 34:54 | ¢ 16-12 32-560]! hl ee we {3 fath. | P — | 20.8.°11 | 17:77 | 34:42 |} ! Ee. » .- 3) 17°94 | 34:42 | — = = ash Ss. = ‘ = $ 18:05 | 34:31 = = = Fr a i | 18-25 | 8429 || — = ie: ee ee. | | + sowieen |(feel7-92 aly eBaz29 pal. See = ¥ “ ue ia xm 21. 8.711 || 16°92 | 34-51 || — ae ae a 2 De nwa 205, “ | 16-68 | 34°67 = a8 fed — |ijises | = | | | 22 fath | br W. 206, * | 16-67 | 34-79 — ||(14-00, | @4588 Resi. | The Bills W.207,/ ,, 16-88 — = = = — (1848 | — i | | | 95 futh. ” W. 208,/; | 16°91 | 34-78 = — = = Fie 34:97 | 126 fath. Clare Island W.210,/ ,, | 16-98 | 3463 | = — SS iqaaete | = a 3 | | 120 fath, | | | | ' | | Clew Bay W.216,) ,, (| 17-74 | 34-42 || — =: = = } = | | | | Clare Island W. 222, | 22. $. 711 | 16-34 34°78 | — — 14:94 34°79 || ( 13-96 34°88 | | | | | {17 ath. er fath. | | | | {| | | Clew Bay W. 228, / 23.8. 11 | 17-25 | 34°67 || = — | a = xt 2 W. 230, H) 16:96 | 34°47 ~ {| = | = = bast a Clare Islund Survey—Marine Algae. 15 9 (8) Tides and Currents——The presence of a good ebb and flow on Clare Island gives a wide intertidal region for shore-collecting. At Westport Quay the rise and fall during spring-tides is about 15 feet, and in April and September 16 feet 6 inches, whilst with neap-tides there is a rise of 12-13 feet. On Clare Island the range is, if anything, greater. Low water at spring-tides is in the middle of the day. In certain spots (i.e. between the islands in Clew Bay and at Darby’s Point, Achill Sound) there is a strong current, and the vegetation is of a different character from that found in other parts. Ballacragher Bay, owing to its narrow opening, has but a small ‘tide—not more than 5-6 feet—except during springs. 2, PHysicAL NATURE OF THE COAST AND CONDITIONS OF EXposuRE, Our district presents great diversities in the physical nature of the shore ; and in describing the same it will be both convenient and natural to divide it into two areas, viz. (1) Clare Island, and (2) the mainland. Speaking generalky, the island presents an exposed. rocky coast, whilst the mainland consists of sheltered ground. In view of the possibility of future work in the neighbourhood, a detailed account of the coast-line is given. (1) Clare Island.— With the exception of the bay near the harbour and a few sandy patches elsewhere, the whole shore is rocky and cliff-bound. Taking the different parts of the island in order, we find that on the south side the rocks are composed of Silurian slate, which forms flat slopes where the beds lie more or less horizontally, but which is very sharp and rugged where the strata are upturned. Rock-pools, narrow gulleys, and deep channels are frequent, and these were found to be useful, as this part of the shore is as a whole somewhat rough and exposed for general collecting, The eastern end of the south shore, and from the corner of the bay to Kinnacorra, is composed of Carboniferous sandstone, a formation found to afford excellent collecting-ground. Various degrees of slope and exposure are presented; and towards Kinnacorra there is a fine series of flat rocks and a number of shallow pools. The Carboniferous sandstone is continued to Portlea, a semi-sheltered bay on the north-east side of the island. A boulder-beach surmounts the upper part of this bay; and the rocks below are flat, with a vegetation somewhat different from that found elsewhere. Further north, towards the lighthouse, the slate is again in evidence; and here caves and vertical surfaces, receiving little direct sunlight, are found, and several shade-loving species appear in the open. Though bounded by cliffs, the shore at this point is easily accessible. The fine stretch of cliff on the north-west side of the island is for the most part precipitous, and the narrow strip of shore below is very exposed and difficult of exploration. B,L,A, PROC,, VOL, XXXI, B 16 15 10 Proceedings of the Royal Trish Academy. In certain spots it is possible to descend to sea-level, which is found to be strewn with large, irregular blocks and boulders. Little seaweed vegetation, however, occurs. On the cliff, colonies of sea-birds provide the conditions which favour a vigorous growth of Prasiola stipitata. With regard to exposure, Beetle Head (or Kinatevdilla) and the shore below the north-west cliff are open to the ocean, and feel the full force of the Atlantic rollers. The south side is open to the sea from the south-west, and is also very exposed; but it is not subjected to quite such boisterous conditions as is the north-west side. Owing to the prevalence of south- westerly gales, the exposure on this shore must, however, be very severe. A bend at the eastern end towards the castle affords some protection, as evidenced by the change in the character of the Fucus growth. The sandy bay near the harbour, which faces east, provides the only sheltered shore on the island, and a few sand-loving species are found at the northern corner. Beyond the bay, the stretch of rocks leading to Kinnacorra is subjected to moderate wave-action, and on the whole the conditions are semi.exposed. At Alnahaskilla (beyond Kinnacorra) greater exposure is met with, owing to the sea coming in from the north; and for the same reason all points along the north-east side are subjected to rough water, though a certain amount of shelter is found in the intervening bays and inlets. In a general way the north-east shore is decidedly more protected than the south, but less so than Kinnacorra. (2) The Mainland.—When investigating the mainland, it was the usual practice to begin at Roonah Point, at the south-west corner of the Ciew Bay, work round to Mulranny on the northern side, and then proceed to Achill Sound and Bellacragher Bay. In describing the ground it will be convenient to follow the same order. : Roonah Quay faces west, and the rocky shore of Silurian slates at the point is subjected to the same exposed conditions as Clare Island. To the south there is a long stretch of boulder-beach, but the presence of rocks in shallow water provides shelter, and there is a certain amount of fairly good collecting-ground. At the point itself, small bays and caves with partial shelter occur. Turning the corner into Clew Bay the exposure rapidly decreases, and between this point and Bartraw there is a series of beaches composed of rocks, boulders, or sand. The surf-loving Nemalion is found as far in as Old Head, where steep rocks and rough water occur for the last time; shallow water then sets in, and there is much sand. When the islands of the bay are reached, the tide recedes for a long distance, and a certain amount of mud begins to appear, but in several spots a luxuriant littoral vegetation exists. Clare Island Survey—Marine Algae. 15 11 Many of the islands provide flat beaches of sand and stone with good collecting-ground at low water. At Annagh Island there is a salt-marsh. The Bunowen river at Louisburgh, the stream at Belclare, and the rivers at Westport and Newport give brackish and fresh-water conditions. In the innermost part of the bay there are wide stretches of mud, and_ the ground is very poor; with the exception of hasty imspections near Westport and one day at Newport, it was but little explored. The northern side consists largely of sand. At Mulranny there are strips of salt-marsh, also fresh-water and brackish streams. The peninsula of COurraun Achill follows, with a rocky shore of Old Red Sandstone and exposed conditions. The floor of Clew Bay is everywhere soft, consisting of sand with stones. In spring and early summer there is a rich and luxuriant sub-littoral vegeta- tion, which in shallower parts may be easily studied from a boat, or hauled up by a hand-dredge. Banks of Lithothamnium occur, and there are numerous Zostera beds. Between the outer islands, channels with a swift current, where the bottom is usually stony, are frequent. Achill Sound, easily reached by train trom Mulranny, provides perfect shelter, and yet fully saline conditions. At low water, the Sound becomes dry, exposing a large stretch of sand. The water enters from both the north and south, and at the end of the ebb-tide the parting of the streams will be observed to take place at a point about a quarter of a mile south of Achill Bridge. The vegetation is for the most part poor, especially in the summer ; but channels provide good ground, and the stones set out by the peasants yield a fine crop of Fucus. Further south (towards Achillbeg) there are swift currents and deeper water, and here a marked increase in the luxuriance of the vegetation is noticeable. On the north side of the bridge there is much mud; but the peat banks flooded by the tide offer some in- teresting types of plant-associations. Bellacragher Bay, a lough-like inlet surrounded by high hills, ditfers from Achill Sound in possessing deep water, and being less tidal. It is connected with the sea (Blacksod Bay) by a narrow opening at the north-west corner. Several streams descend from the hills; and owing to the water only partly escaping on the ebb, the whole bay must be often more or less brackish. Some samples taken at the end of the dry summer of 1911 showed, however, almost normal salinity. - The littoral region is narrow, and composed of stones or rock. The bottom appears to be rocky, since a vigorous belt of Laminaria is disclosed at low water. B2 15 12 Proceedings of the Royal Irish Academy. Il. THE ALGAL VEGETATION. (1) Historica, Resumé or Previous Work. As the present paper forms the first detailed account of the algal associa- tions in any area in the British Isles, a general introductory sketch of the subject may be useful. Kjellmann’s papers on the algal regions and formations of the Murman Sea (77) and the Skager Rack (78), form the starting point of the modern method of studying algal communities. Kjellmann divided the vegetation into three “regions,” and these he further divided into a number of “ forma- tions,” which were characterized by a distinctive vegetative facies, due to the predominance of one or more algae. The main outlines of Kjellmann’s classification have been accepted by subsequent writers, though of late years changes in terminology and great elaboration have been noticeable. One of the earliest attempts to apply Kjellmann’s methods of study to other countries is contained in a paper by Rattray (’85). The paper, which is a biological and ecological account of the algae of the Firth of Forth, is seldom referred to, though it was apparently the first and, until quite recently, the only one of the kind dealing with an area in the British Isles. In other countries, however, this branch of ecology has received more attention. The Norwegian algal vegetation has been studied by Hansteen (92), Gran (93), Boye (94-95), and Kylin (10), and an account of the Faerdese algae was published by Simmons (99). The Greenland marine algae have been thoroughly studied by Rosenvinge, who, after giving two systematic accounts, furnished a paper dealing with the algological communities (99). The marine algae of the Baltic have been investigated by Svedelius (01), who, in his “ Osterjons Hafsalgfiora,” devotes a chapter to the algal regions and formations found under the peculiar conditions presented in that area. The question of defining the algal regions in the non-tidal Baltic had also been dealt with by Reinke (89). he algae of the Faerdes have been described a second time in the, “ Botany of the Faerdes”; and Borgesen’s treatise “The Algae Vegetation ” (’05), which followed his systematic account, is the most important contribution to this branch of algological literature. Kylin, in his algal flora of the west coast of Sweden (’07), deals with associa- tions and other ecological problems at some length. Joubin (’09) described the marine communities at Roscoff, and published an elaborately coloured vegetation-map. The map will be most useful to students working in the locality ; but floristic and ecological data are at present somewhat scanty for Clare Island Survey—Murine Algue. 15 13 comparative purposes. For the Adriatic, we have Tecket’s general account of the algal vegetation of Trieste (’06). The Iceland marine algae are the latest to receive attention from the ecological standpoint, Jénsson’s work (10) forming a useful addition to his systematic papers which appeared previously. Outside Europe, Syedelius’ paper on the periodicity of algae in Ceylon coral- reefs (706), Skottsberg’s work in the Antarctic (06), and Borgesen’s accounts of the algal vegetation of the Danish West Indies (00, ’01, ’11) are apparently the only papers of the kind which have appeared ; but reference should be made to Weber van Bosse (04), and Schiller (?09), which border on the subject. : Turning to the British Isles, with the exception of Rattray’s paper referred to above, no account of the algal communities found on our coasts has appeared, though, as will be seen later, a certain amount of autecology has been done. The numerous and important additions to the British algal flora recorded by Mr. E. M. Holmes and by the late Mr. E. A. L. Batters, have kept our knowledge of the flora well up to date as far as its purely systematic aspect is concerned. Ecological factors, too, were not disregarded, the “ habitat ”’ of a species having been always given a prominent place by British writers; so much so, that in a few cases the specific distinction of similar plants has been strongly advocated on the ground of a difference in habitat. Casual reference to the “Fucus-belt,’ “Coral-banks,” and “ Laminaria-zone,” shows that the more obyious associations were not overlooked. Rattray (85) treats at some length of the size of individuals when gathered from different localities and from different levels; he also deals with various problems connected with the distribution and development of the spores. With regard to algal communities, he recognizes the “regions” of Kjellmann, and describes a number of “areas” (“ formations” of Kjellmann) characterized by the presence of certain dominant species. His “areas,” whether littoral or sub-littoral, are classified almost entirely with reference to vertical distribution, and there is no attempt to connect them with other conditions such as exposure and desiccation. This results at times in a curious association of names in a given zone, and renders the account some- what obscure to anyone not familiar with the exact topography of the locality. The paper is, however, full of facts, and will be of great interest and value to the next investigator of the algal vegetation of the Firth of Forth. In a memoir on Chondrus crispus by Darbishire (’02), the general ecology of the plant is dealt with, and incidentally reference is made to the principal zones of vegetation that occur on the shores of the Isle of Man, Two 15 14 Proceedings of the Royal Trish Academy. interesting ecological papers by Miss S. M. Baker have lately appeared (09, 10), in which experiments, with a view to determine the causes which bring about zonation, are described ; but the general question of algal associations is not touched upon. The ecology of Ulva Laetuca has been discussed by me at ~ some length in a report to the Royal Commission on Sewage Disposal (’11 4). The subject of plant-communities was not in view, but the presence of a distinct mud-formation and an Ulva-association, though not stated in words, was Clearly demonstrated. N. H. Johnson has furnished two notes dealing respectively with the mapping and nomenclature of marine vegetation (lil, 71%). An account of the algal associations of a definite area in the British Isles was thus urgently needed, and in the Clare Island investigations this study was given a prominent place. It will be noted that in several of the works referred to, the systematic account was published first, and the ecological treatise appeared later. This order of investigation is essential. The flora must be thoroughly mastered, before the groupings of the species can be accurately studied. (2) THE AREAS AND THE UNITS OF VEGETATION. A. The Regions. Kjellmann’s term “region” has been generally adopted by algologists ; but the limits of the three vertical divisions included under this term are variously accepted. In the present account they are taken as under :— The littoral region extends from the level of the highest marine vegeta- tion to that of low-water mark at neap-tide. The swb-littoral begins from the lower limit of the last-named, and extends down to the limit of sea- weed vegetation (ie, about 25 fathoms). The clittoral region follows, and being devoid of algal vegetation, is of no importance in the present paper. The reasons for so defining these regions are as follows :—With regard to the upper limit of the littoral region, this, according to Kjellmann, com- menced at the highest tide-level; but, as several writers have pointed out, the algae on exposed coasts often extend far above the-spring-tide limit. A separate region—the supra-littoral—has been suggested by Lorenz. Borgesen, however (’05), showed that no hard-and-fast line can be drawn between the vegetation of the supra-littoral and littoral, and for this reason he did not recognize the area as a definite region. In Clare Island the supra-littoral vegetation, though often found, is of less general occurrence than, for instance, in the Faerdes. It is largely due to the presence of spray (a factor Clare Island Survey— Marine Algae. 15 15 which is regulated by the exposure and he of the rocks), but shade and terrestrial moisture are also important. The associations, moreover, found in this area are the same as those in the upper part of the littoral region, and often merely extensions of them. I therefore agree with Borgesen in not regarding it as a separate region. With regard to the lower limit of the littoral area, on exposed coasts the low-water mark at neap-tide is certainly the natural line of demarkation. At the same time, on a calm day during spring-tides, the Laminaria and Alaria growth may be exposed, and become dry for a short time; but this is obviously exceptional. On sheltered shores the boundary-line is not so clearly marked; and it is difficult to define the beginning of the sub- littoral vegetation. In Clew Bay it is particularly perplexing; and the flatter the shore, the less marked is the boundary. As time did not allow of special work on this point, the lower limit of the littoral region has been made uniform with that on exposed coasts—namely, the ebb-mark at neap-tide. B.—The Formations and Associations. A good deal of confusion exists with regard to the terms ‘ Formation’ and ‘ Association.’ Kjellman first introduced the word ‘ Formation’ into algological literature in his paper on the algological communities of the Skager Rack (78). He describes an algal formation as a small portion of the whole algal vegetation which is distinguished by a characteristic “ Vegetations-Gepriige,” adding that in a general way these portions of the vegetation obtain their characteristic stamp through one or more algae that predominate. Most other writers have followed Kjellmann’s nomenclature. Tt is, however, more in accordance with the terminology now generally employed to use the term ‘association’ for these communities, and to reserve the term ‘formation’ for communities which occur together in a definite type of habitat. The substitution of the former term for the latter was first put forward by Boérgesen (’05), and the name ‘formation’ was at the same time employed by him for “associations when united together under the same or nearly the same ecological conditions.” Examples of Borgesen’s formations are the Chlorophyceae, the Fucaceae, and the Laminaria communities of exposed coasts. Kylin (07) does not follow this nomenclature, but adheres to the term ‘formation’ with the meaning attached to it by Kjellmann. Jonsson (10), on the other hand, follows Borgesen. Borgesen’s use of the term ‘association’ is certainly more correct; but the 15 16 Proceedings of the Royal Irish Academy. larger communities for which he employs the name ‘formation’ appear to represent groups of associations rather than formations. The term ‘forma- tion’ if used to correspond with ‘salt-marsh formation,’ or ‘sand-dune formation’ of phanerogamie botanists, would be more properly applied to the vegetation covering such areas as exposed rocky coasts, or sheltered muddy shores. These might indeed be called ‘subformations’ by some botanists, in which case the whole marine flora would form one formation—the ‘salt-water formation.’ This, however, is hardly justifiable, and there appears to be no reason why the term ‘formation’ should not be applied to the principal types of the marine vegetation, as well as to the sand- dunes or peat-moors that occur above them. In the present state of our knowledge it is difficult to define these marine formations; but notwithstanding this I have ventured to use the term for the vegetation covering the principal types of ground that occur on our shores. The formations proposed are three in number :— 1. The rocky-shore formation. 2. The sand and sandy-mud formation. 3. The salt-marsh formation. All three are represented in the Clare Island area, and in addition two other types are distinguishable, which are dealt with under the heading of— 4, The vegetation of river-mouths. 5. The vegetation of brackish bays. Each of these five types of vegetation consists of a number of associations, and the latter may frequently show natural groupings. But the associations found vary considerably in different localities, being influenced perhaps most largely by the factor of wave-exposure. It should be clearly understood that the classification suggested above makes no pretension to be final. It is possible that the rocky-shore formation will have to be divided into other formations; but, judging from general field-work in many parts of the British Isles, this does not appear very probable. The flora certainly varies with the nature of the rock, but the changes consist in the modification or re-arrangement of the associa- tions, rather than in any profound alteration in the plant-formation. The general plant-covering of all rocky shores (including chalk) appears to be of the same type; and for this reason it is here regarded as a simple formation, though further work will doubtless show many differences in detail, Clave Island Survey— Marine Algae. iba) a The distinction between associations and societies is more puzzling Many of the small communities that have been described by various writers appear to represent societies (as understood by Tansley and Moss) rather than associations; caution, however, is needed, as definite zonation is discernible in one locality, whilst in others the growth is patchy and discon- tinuous. For this reason I have adopted a conservative attitude in the present report, and have used the term ‘association’ instead of replacing it by that of ‘society.’ In the case of communities described for the first time, when the status was doubtful, a non-committal expression has been ‘used. The whole question of transient societies requires careful working out. Many algae appear periodically, and of these some are short-lived, being conspicuous only for a few weeks, whilst others require six or eight months to complete their life-cycle. It is clear, therefore, that before the status of these communities can be settled, a more complete knowledge of the seasonable range of many species is needed, as well as a more detailed analysis of the vegetation. 3.—THE VEGETATION OF THE AREA. i—The Rocky-shore Formation. The rocky-shore formation embraces every form of hard ground, from the exposed rugged slates of Clare Island, to the flat low-lying sandstones of Clew Bay. Great variety in slope and exposure is met with, and also in the surface and hardness of the rock. Speaking generally, the formation is characteristic of open shores. In exposed localities it is the only one to be found. On sheltered shores it is not infrequently poorly developed, or replaced by the sand-and-rock or sandy-mud series. A characteristic feature is the strong basal disk by which the plants are attached, this being true both of exposed and sheltered coasts. Some of the smaller species have, however, other methods of attachment, as explained later. The vegetation of boulder-beaches is included in the formation, and also that of more or less buried rocks and stones, though, in the latter case, the vegetation clearly approaches that of sand-and-rock (pp. 65-67). The pebble-attached association of quiet bays, being characteristic of a soft bottom and not of rock, is placed pro tem. under the sand-formation. The associations differ markedly according to the amount of exposure, and two series are to be recognized—namely, exposed and sheltered. The littoral and sub-littoral vegetation must also be distinguished ; but, except for these, no further sub-division has been attempted. ‘The hardness of the rock, and k,1.A. PRGC., VOL. XXXI- C 15 15 18 Proceedings of the Royal Irish Academy. the degree of slope intluence the vegetation, as does also the nature of the rock-surface ; but, as stated on p.16, this produces merely a change in the associations, rather than a fundamental difference in the plant-covering. Boulders may be singled out for notice, since their vegetation is peculiar and well marked. The associations on a boulder-beach are much broken up, and secondary series on the vertical faces of the rock are of general occurrence (see Plate II). The tops of the boulders are usually bare, owing probably to immediate escape of the water, and to extreme desiccation, which prohibit respectively the retention and development of spores. Points such as these continually present themselves when studying the communities, and several are noted in the report; but it is too early to offer generaliza- tions. A word may be said as to methods of attachment. In addition to the stout disk typical of the formation, three are distinguishable. In the case of simple filamentous species such as Bangia, Urospora, and Ulothrix, each thread is fixed to the rock by the basal cell, with or without the assistance of corticating filaments. Hence the association, which consists of many thousands of minute plants, does not require a rough surface, but occurs also on smooth rocks where other species obtain no footing. It is in reality attached along its whole base. The second type is that of Ceramium acanthonotum, which possesses rhizoids. Such plants cannot grow on smooth rocks, but occur on mussels and barnacles, amongst which they thrust a dense system of rhizoidal filaments. Where Mytilus and Balanus form extensive sheets, these algae are often plentiful. The third type—a soft, spongy disk— is supplied by Callithamnion arbuscula, This is intermediate between the two former and the stout cellular disk. The spongy disk is composed of filaments more or less free at the margins, but forming a dense tissue in the centre. Algae possessing this type require a rough surface, and are commonly found on barnacles. Many transitions exist between the various forms of holdfasts, and the subject, which is of great importance in connexion with the rock-surface, requires detailed study. Epiphytes are adfixed in a number of ways; see Tobler (06), Menz (’10), and Delf (12). In spite of the above exceptions, the ‘majority of the algae forming associations on a rocky shore are attached by a stout. disk (Fucus, Porphyra, Rhodymenia). Clare Island Survey— Marine Algae. 15 19 The following table presents the associations and other communities composing the rocky-shore formation in the Clare Island area :— A. Exposed coast series. (a) Littoral Region. Hildenbrandtia-Verrucaria. Prasiola. Enteromorpha intestinalis. Fucaceae. Lichina. Porphyra. Bangia-Urospora- Ulothrix. Rhodymenia. Laurencia-Gigartina. Callithamnion arbuscula. Nemalion. Himanthalia. Corallina-Lithothamnium. Pool vegetation. Cave vegetation. (b) Sub-littoral region. Laminaria. fncrusting algae. 1B}. Sheltered coast serves. (a) Littoral Region. Hildenbrandtia- Verrucaria. Fucaceae. Lichina. Porphyyra. Laurencia-Gigartina. Corallina-Lithothamnium. Corallina-Cladostephus. Rhbodochorton floridulum. Sand-pool vegetation. Cave vegetation. (b) Sub-littoral region. Laminaria. Encrusting algae. Fruticulose Lithothamnium. A, Eeposed Coast Series. (a) Lnttoral Region. Hildenbrandtia- Verrucaria Association. The description of this algal and lichen community is somewhat brief, owing to the difficulty of distinguishing the species im situ, and the labour involved both in collecting and naming material. Hildenbrandtia prototypus and the marine species of Verrucaria are usually taken together as forming one association, and some writers include other plants such as Rivularia atra and Calothriz scopulorum. It is, however,a question whether it would not be advisable to include in the association all the littoral encrusting species that are soft and not calcareous (see note on sheltered form of association, p. 51). Hildenbrandtia and Verrucaria resemble each other precisely in growth- form ; and they are often found growing together covering an area of several 15 20 Proceedings of the Royal Irish Academy. square yards. Both possess exceedingly thin encrusting thalli, which are so closely adpressed to the rock, that the latter appear as if coated with paint. In addition to those mentioned above, probably several other species of lichens are present. Of the three plants referred to, V. maura forms a band a short distance above the Pelvetia zone, whilst the other two appear below Pelvetia, and descend nearly to the level of low-water. Hildenbrandtia is more susceptible to drying up than V. mucosa, and in the Pelvetia and #. spiralis zones it occurs under the shade of algae, in damp chinks, or in pools. On ordinary open shores its growth does not occur above this level; but on very exposed coasts when there is spray, it ascends far above high-water line, being found in dark fissures in the rock, or in small pools that occur on the rugged cliffs, the water of which is often brackish. On the lower part of the shore Hildenbrandtia is not confined to pools and chinks, but is found on bare rocks, and remains quite dry for several hours. In the lowest levels, it is usually replaced by a broad band of Ralfsia clavata, in which patches of Petrocelis cruenta are ftvequent, especially if the rocks are smooth. Verrucaria mucosa grows mostly on halt-tide rocks that are fully exposed to light and air; in shady positions or where a covering of algae prevents excessive desiccation, the growth extends to the F. spiralis zone; but it rarely, if ever, takes advantage of pools to ascend to a higher level. In this respect it differs from Hildenbrandtia. The plant prefers smooth rocks or boulders, and is seen at its best on the beach north of Portlea, where it covers the upper sides of the boulders from about half-tide level to below the low-water limit of neap-tides. Its growth here is almost pure, and the individual patches are very large, some noted measuring a yard across. The vertical range on the shore varied somewhat in the different localities; but this may possibly have been due to the presence of more than one species. On only one or two occasions was a Verrucaria-growth noted in a rock-pool. Probably few lichens are so difficult to determine or so little understood as the marine Verrucariae; and it is a source of great satisfaction to learn that Miss M. C. Knowles is working at the problem, and doing so from an ecological as well as from a systematic standpoint. The encrusting algae also are in need of critical study. The present community is well developed in all parts of the survey-area. It is abundant on smooth slabs and boulders, and is also found on the rough slate and sandstone. It extends to sheltered localities, where, if rocks and boulders are absent, it often occurs on stones and pebbles. A very similar vegetation exists in Sweden, Iceland, and the Faerdes, according to the accounts of Kylin, Jonsson, and Borgesen, Clare Islund Survey—Murine Algae. 15 21 Prasiola stipitata Association. This association as it occurs in the Survey area is characteristic of bird- cliffs ; but during the winter months it is found to a small extent outside these areas. It is well developed on Caher Island, The Bills, and below the bird colonies near the Signal Tower on Clare Island. The growth is most abundant on the shady side of the cliffs, where it ascends to a considerable height above high-water level, and forms the uppermost association of the marine algae. With P. stipitata occurs, in the upper part, a certain amount of P. crispa var. marina. The localities referred to are very exposed, and; in rough weather, the cliffs are constantly soaked with spray, though during calm periods the vegetation must remain dry for some days. The fact that, in June, the small part of the south side of The Bills that was examined showed no trace of Prasiola, suggests that in summer the growth is confined to the more shady situations—a view which is supported by points noted in the following paragraph. During the winter and spring months, Prasiola is found apart from birds. Wide patches were noted at Roonah and Old Head during the February visit, and in April, 1911, in many spots on Clare Island.. The growth is practically pure, but is sometimes mixed with a certain amount of Calothrix and Lyngbya; it occurs on both smooth and rough rocks. In these localities, as on the bird cliffs, Prasiola forms the uppermost algal vegetation, frequently reaching the Lichina confinis band, but more often forming patches between that zone and Pelvetia. In several spots on the south shore of Clare Island it was present only on rocks sloping landwards, where it was consequently somewhat shaded; but in others it was found on sun-dried rocks and well above the uppermost Pelvetia. These rocks were fully exposed to the surf; and the plants, except during the calmest weather, would be splashed with spray. By July all this vegetation had disappeared; and it was not noted during June of the previous year. The question of the connexion of Prasiola stipitata with bird-colonies is an interesting one. ‘The preference shown by the fresh-water species, P. crispa, for localities soaked with ammoniacal liquid is well known; and its marine relative evidently shares the same avidity for nitrogenous compounds. During a trip to Belfast in connexion with the growth of Ulva in sewage-polluted water, P. stipitata was, on the shores of Belfast Lough during April, noted ‘only around small drainage outfalls. In Larne Lough also a vigorous crop was always to be found in similar situations. At Ballycastle, Co. Antrim, however, as in Clare Island and Clew Bay, numerous patches of Prasiola oceurred (chiefly in shady spots) where there was no special supply of nitrogenous food. 15 22 Proceedings of the Royal Irish Academy. The vegetation described above, taken together with the Enteromorpha growth, corresponds to Bérgesen’s “Chlorophyceae Formation.” In Clare Island, the Enteromorpha vegetation is entirely distinct from Prasiola, and it occurs only where certain definite conditions are present. In the Faerdes, on ~ the other hand, species of these two algal genera are of general occurrence, and form together a well-marked green belt around the islands, The same is the case in Iceland. Enteromorpha intestinalis Association. As remarked above, the Enteromorpha association of exposed coasts is included by Borgesen in his Chlorophyceae formation. In Clare Islands however, the Enteromorpha growth differs so markedly from Bérgesen’s com- munity that it is necessary to regard it as a distinct association. In the Faerdes the formation is described as stretching like a green belt along the coast, and composed of four associations which occur in the following sequence :—Prusiola erispa, Rhizoclonium riparium, EH. intestinalis, and P. stipitata. In Clare Island the Enteromorpha association is quite distinct from that of Prasiola, and is found, not as a continuous band, but only where fresh water exudes from the cliffs. This type of green vegetatiun, though not previously described, is frequently met with in the British Isles. It is not largely developed in the Clare Island district, but is found in its typical form, and is therefore included in the present account. On crossing from Achill Sound to the island, the association in question strikes the eye as a bright green band near high-water mark, between Kinnacorra and the harbour, and wherever a band of this nature is seen it is a sure sign of the presence of fresh water. Where only a small amount exudes, or the shore falls rapidly, the band is narrow, and a fucaceous or other community follows it at once; but with more water and a gradual slope the association takes the form of wide-spreading bands which run transversely over the rocks with a copious development below of Cyanophyceae and Diatoms. A small amount of Rhizoclonium ripartwm is, on flat rocks, sometimes intermixed with Enteromorpha; but more often the growth is practically pure, var. cornucopiae being usually plentiful. On vertical rocks and below small waterfalls Rhizoclonium is more abundant and may be dominant. The forms of Enteromorpha present are mostly short forms of &. intestinalis var. genuina and var. cornucopiae, with the addition occasionally of var. micrococea. With regard to the bathymetric range of the association, its upper limit varies directly with the position and manner in which the fresh water oozes from the rocks. In some places the growth is completely submerged at high tide, but in other spots the upper part is not even reached by the spray of an Clare Island Survey— Marine Algae. 15 23 ordinary spring-tide (e.g., below the lace school on Clare Island), With an alga such as Enteromorpha intestinalis, which is not dependent on salt water, this is only to be expected. By the end of summer much of the growth may be killed through drought, and the lower part of the association then appears as a pure white band of dead tissue. In rock-pools above high-water mark an Enteromorpha vegetation is found on exposed coasts which may be regarded as merely a pool extension of the community just described. The pools in question usually have a large admixture of rain-water, and during the less sunny parts of the year, they contain a vigorous growth of Enteromorpha, and occasionally Hildenbrandtia and Cladophora sericea. Fucaceae Association. (Exposed Coasts.) The conspicuous growth of the Fucaceae that is found on all rocky shores of the British Isles is well represented in the Survey area. The large size of the plants concerned, and the readiness with which the species can be recog- nized, render the Fucaceae association one of the most useful and instructive for ecological study. With few other genera can the effect of wave-action on an exposed coast be studied with so much ease, and with few other associations can the differences in composition and in plant-form he so readily traced. On sheltered coasts the study is more difficult. Though familiar in a general way to all British algologists, no definite account of the Fucaceae associations of our islands has been published, and the study of the vegetation as it exists in Co. Mayo was not without value in bringing to light several interesting features. The composition of the association as it occurs in the British Isles is as follows, the species occurring in the order given from above downwards :—. ® Pelvetia canaliculata. Fucus spiralis. Ascophyllum nodosum (moderate shelter necessary). Fucus vesiculosus. F. serratus. F. ceranoides (admixture of fresh water necessary). An additional species, F. inflatus, was recorded by Borgesen for the Shetland Isles, and Mr. E. M. Holmes has lately received a specimen of this boreal species from Lewis, so possibly it is a regular constituent of the association in the north of Scotland. Besides the ordinary forms and varieties, some very remarkable modifications occasionally occur. (See p. 80.) Fully exposed Coasts,—Turning to the Fucaceae association of Clare Island, 15 24 Proceedings of the Royal Irish Academy. we find that in fully exposed localities it is composed of three species only, namely—Pelvetia, #. spiralis, and F. vesiculosus, var. evesiculosus, a precise combination which has not previously been described, unless it be that referred to by Boye (94-5). Pelvetia forms a narrow zone about 14-2 feet deep. F. spiralis follows with a band only slightly deeper, and composed of fronds seldom more than 6 or 8 inches in length; whilst the lowermost zone of F, vesiculosus, var. evesiculosus, has a greater bathymetric range, and therefore covers a wider horizontal surface. The fronds are very narrow and the growth short, though, except in the most boisterous spots, it is longer than F. spiralis, This type of vegetation is found more or less along the whole south shore of Clare Island, where there is full exposure; and it forms a band covering the entire upper part of the littoral region. The shore is for the most part steep; hence the bands are narrow. With a flatter shore and less exposure, the association spreads out horizontally, and other differences are — noticeable as mentioned later. Of the three species, Pelvetia is perhaps the most susceptible to severe exposure, and F. spiralis the least so. In a general way Pelvetia forms a band just above the ordinary high-water level, but it is splashed daily,except in spells of exceptionally calm weather. In very open spots, or where waves break and there is much spray, it ascends much higher. Everywhere it forms a band that is very conspicuous in summer and autumn by its orange-yellow receptacles. Owing to their bathymetric position, the two species of Fuci, unlike Pelvetia, feel the full force of the waves, but it is only in places with the severest exposure that the association is wanting. The attachment to the rock is exceedingly secure, and, though often torn by the waves, none but ola and worn-out specimens become detached. Several spots were noted where F. spiralis only occurred, and it appeared that the exposure was too great for E. vesiculosus. When fruit is absent it is difficult to distinguish the two species ; but in fertile specimens the short, rounded, hermaphrodite fruits of F, spiralis, and the narrow, pointed, unisexual receptacles of F. vesiculosus separate them at once. On exposed coasts the former plant fruits from spring to late autumn, the latter in winter, spring, and early summer. Moderately exposed Coasts—On passing to less exposed rocks or to bays where there is a measure of shelter, a difference in form and in composition is at once apparent. Pelvetia remains the same, but takes a somewhat lower level. F. spiralis becomes longer and the fronds wider, but still retains its typical spiralis form. F. vesiculosus is also larger, and often possesses a few bladders, the size of the frond and the number of bladders increasing with the amount of shelter. A fourth species, /’. serratus, now enters the association. This begins in the lower part of the F. vesiculosus zone (slightly above Clare Island Survey— Marie Algae. 15 25 Himanthalia), and forms a broad band extending down to the Laminaria level. It appears to enjoy moderate wave-action, but requires flat rocks, and disappears at once where the rocks are steep or the exposure too great. This form of fucaceous growth occurs typically on the slopes near Kinnacorra, near the Castle, and at several spots on the south shore, With greater shelter further differences are evident, Pelvetia and F. serratus remain as before, but F. spiralis broadens out into var. platycarpus and F, vesiculosus develops more vesicles, the length of both species increasing and assuming the heavy bulky growth characteristic of sheltered coasts. The horizontal distribution also of all the species tends to increase, owing to the shore being usually flatter. Such growth occurs in several bays on the island, and is common in more open parts of Clew Bay. Ascophyllum nodosum requires considerable shelter ; and in Clare Island it is present in one spot only, namely Ooghbeg on the south shore. The shore here is rather flat, and the force of the waves is broken by the presence of blocks and boulders. At Portlea (parts of which might appear to be equally sheltered) Ascophyllum is absent, but the presence of a vigorous Himanthalia vegetation indicates greater exposure than might be supposed. For further notes on this species see p. 55. Epiphytes—The typical epiphytes of the exposed Fucus association are as follows, and in the most exposed spots these are practically all that occur :— Ulothrix pseudoflacca (in spring). Elachista fucicola. Enteromorpha compressa. Porphyra umbilicalis. Ectocarpus luteolus. Where the conditions are less boisterous (e.g. at Kinnacorra) a number of other species occur in addition, for example— Ectocarpus tomentosus. Polysiphonia fastigiata (on F. spiralis). E. fasciculatus. Ceramium rubrum. Porphyra leucosticta (in spring). C. Boergesenii. Rhodymenia palmata. Subvegetation.— This, though always better developed in the lower than in the upper levels, is largely dependent on the denseness of the Fucus growth, ie. on the amount of room and light available. When conditions are favour- able, the first four of the following communities are usually represented, and the last three are occasionally present in addition :— Hildenbrandtia-Verrucaria Association. Lichina pygmaea Association. Porphyra umbilicalis Rhodymenia palmata ,, Laurencia pinnatifida $5 Lithothamnium Lenormandi Society. Corallina spp. Society RB, I. A. PROC., VOL. XXXI. D 15 15 26 Proceedings of the Royal Trish Academy. In the low littoral region, if the / serratus vegetation is not dense, a very large number of species occur, the shelter and shade being highly suitable for :— Dictyota dichotoma. Polysiphonia fruticulosa, Chylocladia ovalis. P. thuyoides. C. articulata. Plumaria elegans. Delesseria alata. Lithothamnion polymorphum. Nitophyllum laceratum. Sporelings, many spp. Comparison with other Countries.—In a general way the Fucaceae association described above represents that of the whole of the British Isles, It agrees well with that observed on the south and east of England, and though the Fucaceae of the British Isles have never been specially investigated from the present point of view, it is not likely that any startling departures from the Clare Island type will be met with. The greatest variation may be expected on the north of Scotland. The vegetation described for the Norwegian coast is similar, agreeing floristically, but differmg in minor points. With more thorough investigation some of the latter will probably disappear. In west Sweden, Pelvetia is absent, and Kylin makes no reference to F. spiralis on exposed coasts. He notes the presence of the form #’, Aveschougu on mederately open shores, and it would be remarkable if the short exposed form, so addicted to boisterous localities elsewhere, were absent. A belt of an evesiculate variety of FI vesiculosus is well developed, but from specimens sent by Dr. Kylin, it is clear that his form (compressus racemosus, Kjellman) is different from that which occurs on Clare Island. The Faerdese Fucus vegetation differs in several respects from the Irish. In the first place, I’, serratus is absent, Ff. inflatus taking its place; F. spiralis is not found with extreme shelter, and /. vesiculosus disappears entirely with exposure; lastly, Pelvetia is absent from exposed coasts. The last two points were specially investigated by Dr. Bérgesen during his visit to Clare Island; and he agreed that both species flourish in Ireland with very much more exposure than in the Faerées. The Fucus vegetation of Iceland is similar to that of the last- named, in that it possesses a vigorous growth of Ff. inflatus, though F. serratus is present in addition. From the brief account given by Joubin ('09), the Fucaceae vegetation at Roscoff appears to agree in the main with that of the British Isles. The Lichina communities. Though a lichen and not an alga, Lichina pygmaea is always found between the tide-marks, and forms so essential a part of the marine vegetation that it Clare Island Survey—Marine Algae. 15 27 deserves mention in the present report. The second species, Z. confinis, is scarcely littoral, being confined to rocks just. above high-water line, but its growth may be suitably described together with that of Z. pygmaea. The zones formed by these two species at times approach one another, but they never overlap. Both occur on sheltered as well as on exposed coasts, though for LZ. pygmaea rocks or boulders are essential. With the exception of Joubin (09), algologists have not dealt with this vegetation, though Nylander, as long ago as 1861, drew attention to the belts of Lichina on the sea-shore. (a). Z. pygmaea, the larger and more conspicuous species, forms circular patches which vary from a few inches to a foot in diameter. It usually commences immediately below Pelvetia, and extends downwards through the F. spiralis belt to F. vesiculosus, or to within a few feet of low-water. In certain localities it extends well up into the Pelvetia band; but it never occurs above it. The maximum development usually takes place in the upper part of the zone, where the plant often forms a very conspicuous black band on the vertical faces of rocks or boulders capped with F. spiralis, It undoubtedly favours sloping or vertical surfaces; but it also occurs on those that are horizontal. Though frequent on smooth boulders, it prefers rough or eroded rocks, and is abundant on barnacles. Unlike ZL. conjinis, the present species can only stand a certain amount of drying. In the lower part of the associa- tion it occurs fully open to the sun; but in the uppermost levels it is usually found in chinks and pockets which retain moisture, or on rocks with a shady aspect. In calm weather, during neap-tides, the Pelvetia zone may be left entirely dry for several consecutive days; and any growth of L. pygmaea that reaches this level is occasionally subjected to like conditions. On the exposed and semi-exposed shores of Clare Island and the mainland L. pygmaea is abundant, but where the exposure is excessive it does not appear to flourish. On the large blocks and boulders to the north of Portlea it is one of the few macroscopic plants which form a definite association; and in such localities, where Fuci are absent, the lichens form a useful means of determining tidal levels. A very luxuriant vegetation of Verrucaria spp. covers the boulders of this beach almost from top to bottom (p. 20), During the summer (July-Sept.), a fine growth of Rivularia bullata is frequently found epiphytically on Z. pyymaea. In 1911 it was unusually abundant, being noted in August in all the exposed parts of the mainland and Clare Island. £&. bullata is not confined to the Lichina patches, but also occurs on bare rock, It sets in about half-tide level, and continues to low-water mark. (b). The second spécies, Z. confinis, has fronds barely 3 mm. long, and the individual patches are 2-4 cm. in diameter ; being black in colour it is very D2 15 28 Proceedings of the Royal Irish Academy. inconspicuous on the dark slaty rocks of Clare Island. It usually occurs a short distance above Pelvetia, and extends up the cliffs on the south shore to a height of 12-15 feet. In this position it is frequently drenched with spray, though in summer it may be dry, except for rain, for weeks together. With less exposure its vertical range is restricted. On the gentle slopes at Kinnacorra it is only 3-4 feet above Pelvetia; whilst on the flat shores of Clew Bay and Achill Sound it hardly exceeds the neap-tide level, and is regularly covered during the highest tides. This occasional flooding by spring- tides in sheltered localities obviously takes the place of the soaking by spray on open coasts. In Clew Bay and other places where rocks and boulders are absent, the plant often occurs on flat stones, and may even be traced as an irregular band running along the shore. In Roundstone Bay the three plants, L. confinis, P. canaliculata, and L. pygmaea were noted on large slabs, forming luxuriant and well-marked zones, each species following the other as a band, 1-2 feet deep. The Lichina vegetation at Roscoff described by Joubin (’09) is evidently very similar to that found in Ireland. Both species are present, and the plants show a preference for exposed situations. Porphyra umbilicalis Association. Porphyra is one of those algae which occur as a definite association on both exposed and sheltered shores. The form assumed by the plant is very distinct in the two situations; and it is curious that it has not been seen as an association in localities offering intermediate conditions, although its occurrence in small quantity is common enough. Another feature which gives interest to the exposed Porphyra community is the diversity of its bathymetric range in different countries, and even in different spots in the same locality. For instance, in Clare Island the upper limit of the Porphyra belt is almost always below the Pelvetia zone, whereas in the Faerées the greater part is normally above that belt, and on exposed shores it may occur as much as 40-50 feet above high-water line. Doubtless surf and spray are largely responsible for its presence in such positions, just as they are seen to be directly responsible for minor variations on Clare Island. But this alone does not account for all the differences that have been noted, and further study is needed, On most parts of Clare Island a certain amount of Porphyra vegetation is present; in some it is extensive and forms broad or narrow bands; in others it is limited, and occurs only as small patches between other com- munities. It prefers rocks of a moderate or rather steep slope, but is also Clare Island Survey—Marine Algae. 15 29 found on flat rocks, and rarely on the perpendicular faces. The rock-surface may be rugged or smooth, and a very favourite substratum is that of mussels. In all cases the rocks are exposed; and often the most vigorous patches are those where the waves strike with great force. The normal position of Porphyra is in the upper part of the littoral region, where it occurs as a band a short distance below Pelvetia. In some cases the band is quite narrow, as is shown in Plate I, where it occurs between Pelvetia and F. vesiculosus, taking the place of Fucus spiralis; in others it is mixed with / spiralis, and runs down through the wider belt of F. vestewlosus that follows. Porphyra also occurs where the Fucaceae are absent, and is here commonly mingled with Bangia or Ceramivm acanthonotum. At Kinnacorra it is found on smooth sloping rocks devoid of vegetation, and forms a fairly wide band from a short distance below high-water mark down to about half-tide, at which level Bangia takes its place. In this position, being quite unprotected from the sun, if becomes very dry and disappears during summer. A very distinct aspect is acquired by the association during winter, owing to the presence of an abundant growth of var. linearis. This plant, though often regarded as a distinct species, represents sporelings of P. wmbilicalis, which at that season are elongated and tapering. The growth appears first in October, and develops rapidly during the winter, frequently covering boulders and smooth rocks over extensive areas. It is usually quite distinct from the normal Porphyra association, and forms a band above it. In March the effect of drying winds and increased sunlight is felt, and much of the growth dies. By April var. linearis has disappeared, except in lower and shaded places. With regard to the supra-littoral growth, the contrast between Clare Island and the Faerdes is most marked. In a few spots Porphyra was noted as extending into the Pelvetia band on Clare Island; and at Alnahaskilla it was found at the highest tide-level on the north side of steep bare rocks, a sample of the kind of growth that takes place on a much larger scale on the Faeroes, Below the big cliff on the north side of the island, where the shore is steep and the rocks fully open to the Atlantic, Porphyra was one of the few algae that were noted in the littoral zone. But even here, with shade and continual spray, there was no supra-littoral growth comparable to that described by Borgesen. Its growth probably exceeded the limit of high-water level (which, owing to the constant swell, is not easy to determine), but not to any remark- able extent. In the FaerGes, on the other hand, even in sheltered localities, the lowest iimit is at high-water mark, and in exposed situations it is not found till far above that level. Kjellmann and other Scandinavian alyologists have regarded the Porphyva 15 30 Proceedings of the Royal Irish Academy. community as a winter vegetation; and Borgesen states that, though in the Faerées it is developed all the year round, it is in more southern ‘countries characteristic of winter. On Clare Island this is hardly the case. The Porphyra association is in fine condition in July and August, and abundant even on the south shore (vide Plate I). It is, however, more widely spread in winter, owing partly to the presence of a band of var. /inewris ; but the association in the Clare Island neighbourhood certainly occurs throughout the year. Borgesen also remarks that he did not observe the Porphyra vegetation at North Berwick (Scotland) in July, 1909, but Dr. A. A. Lawson informs me that at St. Andrews (about seventeen miles further north) it was plentiful in July and August, 1910; though in the hot dry summer of 1911 it was much less conspicuous. In the south of England it is a winter and spring vegetation, though in July, 1911, the Porphyra belt was still discernible on exposed rocks at Portland Bill. ‘The exposed Porphyra association is also recorded from Iceland (where it closely resembles that of the Faerées), west Sweden (Kjellmann, ’78, Kylin, ’07), and Norway (Kylin, ’10). To sum up, the Porphyra vegetation of the exposed coasts of Clare Island forms a belt in the littoral zone, at a very much lower level than it does on the Faerdes, and does not, even in shady and very exposed localities, grow far above the high-water line. In contrast to the south of England, Sweden, and Denmark, but in agreement with that described for Norway and the Faerdes, it exists through the entire year. The form of the plant on exposed coasts is entirely different from that found in sheltered localities, Bangia-Urospora-Ulothrix Association. The most striking feature of this community is its sporadic appearance. During the first two years of the Survey scarcely a thread of Bangia or Urospora was found; but during the spring of 1911 a remarkably fine and widespread development of the typical Bangia-Urospora-Ulothrix association occurred on Clare Island and on the exposed parts of the mainland. This vegetation was probably at its best during the time of the April visit; by August it had entirely vanished. In the Faerées it appears to be more or less permanent; but, as shown below, its sporadic appearance is known in other countries, and it may even be absent for several years in succession over a wide stretch of coast-line (vide Borgesen, 05, p. 720). For this reason the appearance of well-marked belts of Bangia and Urospora during the last year of the Survey was particularly gratifying. The community in question occurs on exposed and moderately exposed rocks, especially where the surface is smooth and where other algae do not ootain a footing. In bathymetric range the association as a whole corresponds Clare Island Survey— Marine Algae. 15 31 to that of Porphyra, extending from below the Pelvetia zone to within a few feet of low water. Its composition is as follows :— Bangia fuscopurpurea, _ Ulothrix pseudoflacca, ) co-dominant. Urospora mirabilis, Ulothrix consociata, Enteromorpha minima. As a rule all the species, except #. minima, which is local, occur together, though each forms a separate patch or narrow strip within the association. Occasionally, however, the species occur separately; and a pure growth of Bangia or Urospora may cover a wide area. In a few cases the growth may be thoroughly mixed. - An effort was made, when the full series was present, to determine the vertical sequence of the species; but this was unsuccessful in so far as obtaining a constancy in succession was concerned, The necessity of microscopic examination made the determination in the field difficult, and the work required more time than was available. As a general rule, however, where the association is typically developed, Bangia appears to have a wider yertical distribution than the other species, as it is often both the first to appear and last to leave on passing from top to bottom of the area. But occasionally a Ulothrix growth is found at a very high level in sheltered localities where Bangia is absent. The occurrence of the Bangia-Urospora association on smooth rocks and boulders where other vegetation is absent is explained by the exceedingly fine unbranched filaments of the plants, reducing to a minimum the strain on the basal attachment. At the same time each filament is separately fixed to the rock, so that the association may be said to be attached along its whole base. The most usual extraneous species is Porphyra wmbilicalis, small specimens of which frequently occur in the upper levels; and it is into the Porphyra association that the present community most frequently merges. Bangia, which is less restricted than Ulothrix or Urospora, also occurs on barnacles, where it enters into competition with Ceramium acanthonotum. The total absence of Bangia in some years has been referred to by Boérgesen for Norway (’05, p. 720), and Rosenvinge for Denmark (09). The latter, who has studied the Bangiales of Denmark in detail, writes as follows concerning B. fuscopurpurea :— “The most dangerous condition for the Bangia vegetation is a fairly long period of easterly winds so light that this vegetation is not reached by the 15 32 Proceedings of the Royal Trish Academy. waves, especially when the weather at the time is bright and dry. Its occurrence is therefore very different, not onlyat various seasons, but in different years. In winter it is very abundant: but the critical period of the spring will every year kill a greater part of it; and on the duration and intensity — of this period depends to what degree that will take place. In summer, for example, it occurs at Frederikshavn in some years only in small quantity, while in others it forms extensive growths—as in the beginning of August, 1902” (Le., p. 58). With regard to Clare Island, drying winds during a critical period of the life-history no doubt largely influence the growth of the plants. The association may occur annually as winter vegetation, and it may possibly have been present in the early spring of 1910; but if this was the case, it had completely disappeared by June, and was certainly absent in April, 1909. Comparison with other Countries.—An association corresponding to that described has been noted in many countries. It is present in Norway, Sweden, Greenland, Iceland, and the Faerdes, having been dealt with by Kylin, Rosenvinge, Jonsson, and Borgesen respectively; and, as already shown, it occurs in Denmark. ‘The belt in these countries, as would be expected, is found at a higher level than in Clare Island. It reaches, or extends above, the high-water line in each case; and in the exposed part of the Faerées it occurs as much as 30 feet above this level. In Greenland the association differs floristically, and is termed by Rosenvinge the Monostroma grocnlandicum association, after the dominant species. In the Baltic (Gotland) Svedelius has described a similar vegetation, consisting entirely of Urospora penieilliformis. 3 Rhodymenia Association. An association of Rhodymenia similar to that described by Borgesen, and noted by him as being widely spread on the Faerdes, occurs in our district, though only to a limited extent. Borgesen stated that he had found no reference to such an association, but that he expected it to be common in the North Atlantic. In the Clare Island area it is but feebly developed ; but at Dog’s Bay, near Roundstone, a very extensive growth was observed, showing that the association exists on some parts of the west of Ireland, as indeed was already known. On the island, patches of Rhodymenia occur in several spots between Portlea and the lighthouse, and on the mainland at Roonah and at Old Head. Judging by the localities in which the plant was found, Rhodymenia prefers a northern aspect, which would account for the complete absence of its growth, as an association, on the south shore of Clare Island, At Old Clare Islund Survey— Marine Algae. 15 33 Head it is found on flat, sloping, or vertical, semi-exposed xocks, which face north or north-east. At Portlea it affects similar situations, but shows-a decided preference for vertical rocks (these being at the sametime the more shaded). The plants are attached by a strong basal disk, and are usually found on mussels, but they occur also on limpets-and bare rock. The vertical area covered by the alga is rather extensive, reaching at times from the bottom of the F. spiralis zone down to about low-water mark, but more often forming a band below the upper Porphyra. It is frequently interrupted by patches of other vegetation. At Old Head, Fucus vesiculosus, var. evesiculosus is the chief intruder; but portions of the Porphyra and Callithamnion arbuscula associa- tions above, and the Nemalion vegetation below, are also interspersed amongst it. A modified form of the association occurs on boulder-beaches, the plant hanging in profusion from the lower part of boulders surmounted by Himanthalia (Plate IT). : The form of the alga when growing as an association on exposed coasts is very distinct. Short and tufted, not more than 5 to 4 inches long, it consists of narrow or rounded fronds, with a number of small lateral proliferations. Though narrow, it is very distinct from var. sarniense, which is thinner and paler in colour, and resembles Gracilaria multipartita in form, With increased shelter the plant becomes larger, till at length the pool-form, with fronds a foot or more in length, is reached. ‘The islanders distinguish the larger and small forms as Dillisk and Crannogh respectively, and always associate the latter with the presence of “small shells” —iLe., mussels (see Economic Section), Borgesen remarks that the Rhodymenia association prefers places where fresh water oozes from the rocks. This is not the case on Clare Island, Asa pool or streamlet plant, Rhodymenia enjoysan admixture of fresh water; but when growing as above described, it was always found in a purely marine habitat. Laurencia-Gigartina Association, Near the low-water line on exposed coasts there often exists a dense earpet composed of the two red algae,—Laurencia pinnatifida and Gigartina mamillosa. Gigartina occupies the lower part and goes down into the sub- littoral region, whilst Laurencia extends more or less in an upward direction. The band formed by the latter is often conspicuous, so much so that it was at first regarded as a distinct association. Further examination at spring: tides showed that it was frequently connected with a zone of Gigartina, and that in the more exposed places the latter species was dominant. It gradually became evident that their relative abundance was largely a matter of exposure and slope, and that these two species of similar habit could ‘be | B,I,4. PROC,, VOL. XXXI, a0) 15 15 34 Proceedings of the Royal Trish Academy. naturally regarded as forming a single belt. In a general way Laurencia prefers a certain amount of shelter, and thrives on sloping or flat rocks, whereas Gigartina enjoys the roughest water and steep surfaces. On comparing this plant-community with those of other countries, it was found that the joint belts formed by the two species in Clare Island corresponded to Boérgesen’s Gigartina association in the Faerées—a conclusion which was confirmed by him during his visit. It was thought best, therefore, to connect the Irish type of vegetation with the Faeréese, and the name Laurencia-Gigartina association has been given. In the Faerdes Laurencia is very rare, being found only in pools. The Gigartina band may be dealt with first. On exposed coasts it is barely uncovered during neap-tides, but with spring-tides a distinct belt reaching down to the Alaria and Z. digitata zone is revealed. It is frequently accompanied by a certain amount of the narrow form of Chondrus crispus. Well seen on vertical rocks, where it forms a band 2 feet deep, it also occurs on flat and sloping surfaces, here usually as an undergrowth to Himanthalia. On the open coasts it is often accompanied by the following algae, all of . which are known to enjoy the turbulent conditions of low-water mark :— Scytosiphon lomentarius. Plocamium coccineum. Rhizoclonium implexum. Polysiphonia thuyoides. Chylocladia ovalis. P. Brodiaei. Lomentaria clavellosa. Corallina squamata. The Gigartina belt occurs also on the more sheltered ground at Portlea. Tt first appears well up in the Laurencia band; but as the lower levels of the littoral region are reached, it increases greatly in quantity, till finally it occurs as a pure growth covering comparatively large areas. At Kinnacorra, on the other hand, where there are varying degrees of exposure, the Gigartina vegetation is practically absent. Passing now to Laurencia, we find this is more conspicuous on account of its truly littoral habit. It begins shortly below the half-tide level, and runs down into the Himanthalia association. It is common on moderately exposed shores, and appears to prefer clean, gently sloping rocks where sand is absent. Under these conditions Laurencia may exist as a pure growth 2 or 3 yards wide, but where the shore is flat it extends over a much wider area, being then mixed with other species. When sand is present it gradually gives place to Rhodochorton floridulum. In its lower levels the Laurencia belt forms an undergrowth to Himan- thalia, and on sheltered shores occurs in a similar manner beneath F. serratus. It is found at times under F, vesiculosus, and thus runs upwards considerably Clare Island Survey—Marine Algae. 15 35 higher than on bare rocks. If the shade be not too dense, its growth is quite as luxuriant as when fully exposed to light and air. Another species, Z. hybrida, is a characteristic element in the upper part of the association. This plant is more restricted in its range, and appears to be more sensitive to desiccation. Ona flat shore it reaches its maximum development somewhere about the middle of the Z. pinnatifida band, where it is at times co-dominant. In the lowest parts it is practically absent; and in the upper it is found only in the damper positions. On very exposed shores L. hybrida is scarce (see p. 136). L. pinnatifida reaches its maximum development about April or May ; and at this season sloping rocks are carpeted with a growth 4 to 6 inches high. Aiter May many of the larger fronds disappear, and the association as a whole becomes inconspicuous. There remain, however, numerous small shoots, which are continually augmented during the summer, the plant being apparently perennial. JZ. hybrida, on the other hand, is an annual. Young sporelings 1 to 2mm. long are discernible in September ; they develop during winter, fruit in spring, and disappear in May. Very marked colour-changes due to illumination are shown by Z. pinna- tifida, and to a less extent by Gigartina. The normal colour of the former is a deep purplish red; but during summer the growth in sunny situations is pale yellow green. This gradually gives place to purplish green in autumn, which becomes redder as the days shorten. The production of antheridia by Laurencia in March and April gives the plants a golden hue, owing mainly to the yellow substance found in the antheridial cavities. As a large number of male plants are often found together, sheets of bright yellow fronds are frequent. Callithamnion arbuscula Association. The plant-community formed either by Callithamnion arbuscula, ot Ceramium acanthonotum, or a mixture of both, has been described by several writers under various names. Callithamnion is the more conspicuous plant, and most writers have dealt largely with this species; Ceramtwm acanthonotum is, however, an equally important element. Clare Island is peculiar in possessing in addition a small quantity of the southern Callithamnion granulatum. The association is well developed on many parts of the island and exposed mainland. The relative amount of the two species varied during the different seasons—a fact which probably explains discrepancies in accounts of the same area by different writers. In July, 1910, Ceramiwm acanthonotwm was the dominant species on Clare Island, whereas in August, 1911, K2 15 36 Proceedings of the Royal Irish Academy. Callithamnion arbuscula was more abundant. Apart from these seasonal variations, Ceramium is found chiefly on bare slopes with much or moderate exposure; sometimes it occurs pure, forming a dense carpet several yards square, but more often it is mixed and interrupted. Callithamnion prefers the more boisterous positions, and is especially luxuriant on steep or vertical rocks. The belt formed by the association is at the same level as the lower part of the Bangia and Porphyra associations, or slightly higher than the Nemalion and Corallina communities. It usually begins just below F. spiradis, and extends to within a few feet of low-water mark. Callithamnion often appears a short distance above Ceramium, but for the most part the two species occupy the same level. Where fresh-water streams exist, Ceramium not infrequently ascends the shore, and may reach as high as Pelvetia. C. arbuscula, on the other hand, does not thrive in fresh water. Unlike the Bangia-Urospora association, the present plant-community is most frequent on rough surfaces offering a good foothold. Ceramium, which forms a spreading growth, usually occurs on mussels and Corallina, amongst which it pushes a mass of rhizoids; but it also grows epiphytically on the Callithamnion. The latter, on the contrary, has a scattered habit, each plant possessing a single stem, which is thick and spongy. It is most frequently attached to limpets, mussels, or the rock between them; only varely is it found on other algae. Although both species occupy a fairly high level, owing to the retention of water by their spongy thalli neither becomes dry when the tide is out. The associations into which the Callithamnion arbuscula vegetation most frequently merges are those of Bangia, Nemalion, Corallina; and in addition to algae derived from these sources the following species frequently occur scattered in it :— Rhizoclonium tortuosum. Polysiphonia macrocarpa. Enteromorpha compressa. P. Brodiaei. Scytosiphon lomentarius. P. thuyoides. Petrospongium Berkeleyi. Comparison with other Localities—In the British Isles Borgesen has noted the community in Shetland, and I have observed it at. Whitby (Yorks). On the south coast of England, where C. arbuscula is absent, the association is represented by a belt composed entirely of C. granulatum. (= C. spongiosum, Harv.). This has been noted at Swanage and Portland, and is doubtless common. Ceramium acanthonotuwm, though known on the south coast, does not enter into the Callithamnion association in either of these localities. Clare Island Survey— Marine Algae. 15 37 It is interesting to note that, though C. arbuscula was the essential species in Clare Island, specimens of the southern C. granulatum were scattered here and there amongst it. The latter agreed in form with the Dorset specimens, being short, very densely branched, and of a peculiar pale colour—a feature which gives the clue to its presence (see p. 142). Hansteen, Boye, and Kylin have each dealt with a vegetation on tite west coast of Norway that is probably very similar to that here described, though it is Callithamnion that has mostly attracted attention. Kleen also records both algae from Nordland. In the Faerdes, Simons refers to a “©. acanthonotum formation,’ not observing the presence of C. arbuscula, whilst Bérgesen, exploring the coast more thoroughly, found the latter abundantly in exposed places, and adopted the older term. — Both species are found in Iceland, though Jonsson does not allude to them as forming an association. : Some remarks by Harvey with reference to this vegetation may be quoted here. He observes with regard to C. arbuscula that “it delights in the most exposed rocks and the roughest water, and very commonly grows on the shells of Mytilus rugosus” (Phye. Brit., pl. 274); and concerning C. spongiosum, he writes, “It is curious that it appears to occupy the place of C. arbuscula on shores where the latter is not found, these plants never growing together, though both affect similar situations on different shores” (1. ¢., pl. 125). As shown above, Harvey’s statement as to position and habitat of the species holds good for many countries, though the Clare Island work proves that the two species may occasionally occur together. _ Nemalion Association. The present vegetation, described by Kjellmann in 1875, was one of the first algal associations to be recognized. It differs from many others in being a purely summer growth, and might therefore be more suitably regarded as a plant-society. The Nemalion belt occurs in the lower half of the littoral region—that part of the shore which is crowded with a succession of algae, and in which many transient societies are recognisable. Nemalion prefers bare rocks, and usually develops in spots devoid of other vegeta- tion. It would prone yi find a place as a subdivision of Borgesen’s “Corallina formation”; but in the present state of our knowledge ‘it appears advisable to follow Biewous writers rather than introduce doubtful alterations. The Nemalion growth is first noticeable early in June, reaches its maximum in August or September, and disappears in October. The association in Clare 15 38 Proceedings of the Royal Trish Academy. Island (as in 8. England) is composed of two species—JV. multifidwm, and the stouter unbranched JV. elminthoides Batters (= NV. lubricum J. Ag.).’ The zone formed is 5 to 6 feet deep, and extends from about half-tide level to the ordinary low-water mark. Its luxuriance varies considerably in different localities. In some, a vigorous growth is present in the upper part of the zone, whilst in others the algae are better developed near low-water mark. JV. elminthoides is usually confined to the upper portion, and lasts later in the season than N. multifida. The plants are attached to bare rock, or limpets, and occasionally to mussels. They prefer sloping surfaces, fully exposed to waves and surf. The Nemalion association is found in the band of Lithophyllwm imerustans ; but it prefers bare rock devoid of other vegetation. Where mussels occur, it is usually replaced by the association of Ceramiuwm acanthonotwm, or Rhodymenia. Amongst it are scattered individuals of Scytosiphon lomentarius, Enteromorpha compressa, Rhizoclonium implecum, and Polysiphoia Brodiaer, the last-named being confined to the lowest levels. On Clare Island the association is of frequent occurrence; and in Clew Bay it occurs on exposed points, such as Old Head. A similar type of growth has been described by Kjellmann (’78), and Kylin (’07), for west Sweden ; by Gran (’93) for Norway ; it would appear also to be widespread in Denmark (vide Rosenvinge, ’09). In the south of England the community is frequently met with, having been noted by me on the coasts of Dorset, Devon, and Cornwall. Himanthalia Association. This familiar association, which is well developed on Clare Island, is characteristic of moderately exposed shores. It forms a belt in the lowest part of the littoral region; but the extent of its horizontal range varies with the amount of slope and the nature of the coast. On steep and vertical rocks it is absent, or exists as a narrow band, 1 foot deep. On a shore of moderate slope the band is wider, and extends from the Laminaria zone to 2 or 3 feet above low-water mark; whilst on flat shores, especially where there is a dense sub-vegetation, it covers a wide area. With partial shelter, Himanthalia descends some distance into the sub-littoral region, and occurs mixed with Cystoseira ericoides and species of Laminaria; but apart from the extension both in an upward and downward direction, its normal position 1 This plant, clearly figured by Velley, is usually regarded as a distinct species ; but as seen in Clare Island and in the South of England, it is difficult to separate from WV. multifidum. Clare Island Survey—Marine Algae 15 39 on a sloping shore is between the Lawrenciw pinnatyida belt and the Laminaria association. In the form of a narrow band, the Himanthalia vegetation is seen on the south shore of Clare Island in spots where the rocks are steep, and there is full exposure to the waves; as a wider zone, it shows well on rocks which form large sloping slabs, as at Kinnacorra. At Portlea a more extensive growth is met with. The shore here is flat, but there is a moderate amount of rough water and surf. A dense undergrowth of Lawrencia pinnatifida and other algae covers the lower part of the littoral region, and on this the Himanthalia lies as a thin covering. The belt, however, is not so deep as it appears; for, although during spring-tides it is left dry for three or four hours, during neap-tides the time of exposure and the amount uncovered are small. The undergrowth conserves moisture, and enables the alga to ascend to a higher level than otherwise. Portlea also affords an example of another type of growth, viz., that on blocks and boulders. In this case Himanthalia is attached to the upper parts of the boulders; but the strap-like fructifications hang down over the sides (see Pl. II). Here again the alga may ascend above its normal limit owing to the shade between the boulders. Boérgesen remarks that, according to Boye, Himanthalia avoids localities directly exposed to the surf, which is in contrast to its habit in the Faeroes. In the British Isles it is certainly a surf-loving species; and in Clare Island it occurs along the south shore, and also on such exposed points as Alnahaskilla. At Ooghbeg, the one spot on Clare Island where Ascophyllum exists, Himanthalia is found where the waves break at low-water; but as a rule, if there is sufficient shelter for Ascophyllum, Himanthalia is absent. As noted by Bérgesen, the present alga cannot stand desiccation ; and though odd plants are seen some half-way up the shore, they do not thrive unless protected from the sun. As a pool-plant it is one of the commonest species, and is then usually infested by Hetocarpus fasciculatus, Ceramium Boergesenti, and Herponema velutina. In the open, Hlachista seutulata is the usual epiphyte. Corallina and Lithothamnium Associations. A belt of Corallina spp. is found just above low-water mark in many parts of Clare Island. It is characteristic of very exposed rocks, where it appears to take the place of the Lawrencia pinnatifida band of less open spots. An extensive growth of encrusting algae is usually met with as an under- growth, and the two types of vegetation appear to be intimately connected, and may for the present he treated together, 15 40 Proceedings of the Royal Irish Academy, -. ‘The species concerned are as follows :— Corallina squamata, dom. C. officinalis, co- or sub-dom. Lithothamnium Lenormandi. inal Undergrowth, Lithyphyllum incrustans. : J | in zones, Lithothamnium polymorphum., (a) The Corallina growth is very much worn down by the waves, so that it occurs in the form of dense cushions, the specific identity of which is difficult to ascertain. In shallow depressions or with slight shelter, the production of larger and more perfect fronds allows their true nature to be determined, and in each case examined it was evident that the bulk of the growth was composed of C. sguamata, though C. officinalis was at times also present. The first signs of the Corallina belt are found at about half-tide level, usually just below the top of the Nemalion zone. The plants are found here in chinks and fissures, where there is little risk of drying, and on passing downwards they gradually spread out over the surface of the rocks. They are, especially in the upper parts, frequently attached to mussels; and the growth of these two organisms together form the compact cushions referred to. At the lower part of the zone the fronds are large. This is due partly to their being protected by the Laminaria foliage, and partly to their being below the level at which the waves break. A great improvement in colour is at the same time noticeable. C. squamata as well as C. officinalis occurs at the top of the Laminaria belt; and the latter certainly, if not indeed both, descends some distance into the sub-littoral region. With increased shelter, though the plants are larger and less broken by the waves, the band as a whole is not so conspicuous, owing to its being replaced by other algae (usually Laurencia). Shallow pools are an exception. Here a fine growth of C. officinalis is often present, and less frequently C. squamata, ‘The latter is seen at its best on narrow ledges, or hanging from the sides of shady rocks. The Corallina association above dealt with differs floristically from all others hitherto described, in being composed largely of the southern species C. squamata. his is probably the case in all the warmer parts of the British Isles. ‘There is no question as to the abundance of C. sguamata in Clare Island ; and it is doubtless equally common along the whole of the south and west of Ireland. It also occurs in plenty in the south of England, and has been noted as forming the principal constituent of the Corallina association at the end of Portland Bill. (b) The Lithothamnium vegetation is described here together with that of Corallina, because it is exceptionally well developed when found in company Clare Island Survey Marine Algue. 15 41 with a widespread growth of that alga. A thorough study of its occurrence in varying conditions was not made; consequently the following notes must be regarded as but a contribution to the study. The growth in question occurs in three zones, Lithothamnium Lenormandt, the uppermost, Lithophyllum inerustans, the second, and the .third, LTithothamnium polymorphum, appears only when low-water mark is reached. All three plants resent desiccation, and cannot bear being dried up. L. polymorphum, and, to a lesser degree, L. Lenormundi, prefer shade ; but LZ. cerustans thrives in the bright sunshine of shallow rock-pools. The zone of LZ, Lenormandi occurs on damp rocks slightly above half-tide level, at which height moisture is local, being found in fissures, on shady vertical surfaces, or under the foliage of the larger algae. The zone is more extensive than at first sight appears, and is probably widespread in the British Isles. L. incrustans follows. This is abundant in all the rock-pools, and is by far the commonest species in the district. Apart from its growth in pools, L. wnerustans first appears in chinks and hollows in the lower half of the littoral region. As one descends the shore the crusts become larger and more widely spread; and on reaching the Himanthalia level they form large sheets, filling up interstices between limpets, barnacles, and Ral/sia clavata. At low-water mark the plant is joined by ZL. polymorphum, the red warty thallus of which is at once distinguishable from the paler L. inerustans, though the latter at this level assumes its true matve-pink colour, in contrast to its almost white appearance higher up. JZ. polymorphwm extends into the sub-littoral region; Z. incrustans probably does the same, but its growth here on exposed shores was not investigated. The Lithothamnium vegetation described above is specially characteristic of exposed, somewhat bare rocks, where algae such as Laurencia and Himanthalia are poorly developed, but where there is a large growth of the short form of Corallina. In localities where there is less exposure, or where shrubby ‘and foliaceous algae occur in plenty, the Lithothamnium vegetation is poor: some traces of the LZ. Lenormandi and L. polymorphum zones may be found, but L. incrustans usually disappears altogether. Vegetation of Rock-pools. Several different types of pools may be distinguished according to their elevation, and to the geological nature of the shore. A number of other factors must also be taken into consideration—namely, the size, depth, and configu- ration of the pools, and conditions of temperature, illumination, and drainage. Whilst much of the pool-vegetation is perennial and more or less constant, a large part is transient and varies considerably, not only from month to RB. I. A. PROC., VOL. XXXI. FB 15 15 42 Proceedings of the Royal Irish Academy. month, but from one year to another. A detailed study and analysis of the vegetation was impossible; but a series of observations was made, and a general account is given below. In the upper pools an encrusting vegetation is always conspicuous. At the highest levels Hildenbrandtia prototypus is tound, especially on rugged rocks and on exposed cliffs. Slightly below, and generally in flatter areas, Ra/fsia verrucosa appears, covering the bottom of shallow pools with a thick crust, and often completely ousting Hildenbrandtia. At a lower level (probably that of ordinary high-tide), Zithophyllum incrustans begins to develop, first as avery thin crust which does not fruit; then rapidly increasing in vigour on the lower rocks, it passes into the Lithothamnium association (p. 41). Ralfsia and Lithophyllum are, in the upper shallow pools, in competition. The former cakes off with age, allowing Lithophyllum for a time to get the upper hand. Ralfsia, however, tolerates more fresh water, and probably grows more rapidly. Both can withstand intense insolation, and often form large sheets without any protecting vegetation whatever. The water of these pools becomes in summer very warm. With regard to the upright vegetation, Hnteromorpha intestinalis is often alone in the highest pools, but is joined by Cladophora sericea, Chaetomorpha aerea, and Seytosiphon lomentarius, where the salt-water is constantly renewed by spray and splashing. Below these a series of fairly characteristic pools, situated about high-water level, is reached, which are well supplied with seaweeds during winter and spring. But in summer the larger algae are scarce, or represented by a few hardy species, usually infested with Diatoms and Cyanophyceae. The dominant species are :— Cladophora rupestris. Gigartina mamillosa. C. sericea. Laurencia hybrida (except summer). Scytosiphon lomentarius. Polysiphonia fruticulosa. Leathesia difformis. Dumontia filiformis (except summer). Gelidium sp. Corallina officinalis. The additional winter and spring flora need not be listed. The pools are not always replenished in calm weather, and are distinctly brackish ; they are about 6 inches deep, and commonly run parallel with the shore. ‘At alower level the typical purely marine rock-pools commence, in varied form, and with abundant vegetation. They are much more difficult to classify, and have a constantly changing flora. With a sufficient depth, large algae, such as Halidrys, Cystoseira fibrosa, &c., are plentiful, and provide shade; hence almost any of the shade-loving Florideae (except those which require daily exposure to the air) may be met with. If shallow, the flora is more limited, Clare Island Survey— Marine Algae. 15 43 though sporelings and young plants of a very large number of species occur, In Vancouver, Henkel (06) has worked on the structure of rock-pools; and Skinner’s paper (03) is one of the only attempts to analyse the flora and the conditions obtaining. His conclusions, equally applicable to the British coasts, are as follows:—In general, the higher the elevation of the pool, and the less exposed to wave-action, the fewer the species found, though the number of individuals may be great. The more gradual the slope, and the rougher and more irregular the sides, the more abundant the plant-life. A perpendicular or receding wall is unfavourable for the location of plants. An attempt is made below to indicate the characteristic species of ordinary rock-pools not more than 18 inches deep. A complete list is out of the question. In a general way they are at their best in early spring. During the dark days of winter many of the Florideae, which at other seasons thrive in more shady spots, are found in shallow pools (e.g. Phyllophora rubens, Plocamium coccineum, Laurencia obtusa, Rhodomela subfusca, Halurus equisetifolius). Hence in winter and spring the Florideae are markedly dominant. In March and April many transient species develop ; and in May and June brown algae are abundant, and also species of Cladophora. The Florideae in shallow pools are in summer of poor colour; and a great decrease in the number of species is noticeable. Ulva and Enteromorpha spp. may be dominant in August and September. Excluding large species, such as Fucus, Laminaria, &c., and small epiphytes like Myrionema, Elachista, and Chantransia, the most typical species are as indicated in the list on pp. 44, 45. Cave- Vegetation. As would be expected from the conditions obtaining in caves, shade-loving littoral plants abound; whilst in addition species from the sub-littoral region also occur. Provided wave-action is not too severe, the cave-flora is often full of interest biologically, long tunnel-like caves, with pools and boulders, being particularly instructive. Though the Clare Island caves are neither numerous nor extensive, their flora is characteristic and worthy of record. ‘They may be roughly divided into two sets, namely, exposed and: sheltered. The flora of caves fully exposed to the sea is usually poor, incrusting species on the walls being at times the only vegetation present; but sheltered caves, especially if stocked with boulders, are rich in species, and possess the shaggy growth of which Plumaria is typical, as well as the mossy and incrusting coverings supplied respectively by Rhodochorton Rothvi and Lithothamnium polymorphum, Characteristic Vegetation of Rock-pools. Feb., 1911. | April, 1911. July, 1909. | Roonah Pt. |Clare Island. Clare Island. Uct., 1910. Clare Island. Monostroma Grevillei, Enteromorpha clathrata, . E. compressa, Ulva Lactuca, Rhizoclonium implexum, . Cladophora pellucida, C. rupestris, C. sericea, C. glaucescens, . C. albida, a C. lanosa, . Codium adhaerens, C. tomentosum, C, mucronatum, Dictyosiphon foeniculaceus, Litosiphon pusillum, Phloeospora brachiatum, Scytosiphon lomentarius, Ectocarpus confervoides, E. fasciculatus, E. granulosus, Pylaiella litoralis, Ismophloea sphaerophora, . Myriotrichia claviformis, Sphacelaria cirrhosa, Cladostephus verticillatus, . Chordaria flagelliformis, Mesogloia vermiculata, Castagnea virescens, . Leathesia tuberiformis, Dictyota dichotoma, . Porphyra leucosticta, P. umbilicalis, . Pterocladia capillacea, Gelidium pulchellum, G. latifolium, Chondrus crispus, Gigartina mamillosus, Phyllopkora rubens, ie, 8 KX EK ES OE Kee ta ee a eo [eee SS KEES CS SC Eh x X | Dee ON OK, KKK EK LK X a ab. x X X X [exe =| SESS SORES SG OK I OK KR OE KSI || AMS NIN suk Ki JDK ab. ab. EK OX Ok DK Clure Island Survey—Marine Algae. 15 45 Phyllophora membranifolia, Cystoclonium purpurascens, Calliblepharis jubata, Rhodymenia palmata, Lomentaria articulata, L. clayellosa, Chylocladia kaliformis, C. oyalis, Plocamium coccineum, Nitophyllum punctatum, N. laceratum, Delesseria sanguinea, D. ruscifolia, D. hypoglossum, Rhodomela subfusca, Laurencia obtusa, L. hybrida, L. pinnatifida, . fs Polysiphonia urceolata, P. nigrescens, P. Brodiaei, P. fruticulosa, . Pterosiphonia thuyoides, Dasya arbuscula, Spermothamnion ‘Turneni, . Griffithsia setacea, Halurus equisetifolius, Monospora pedicellata, Plumaria elegans, Ceramium rubrum, C. Boergesenii, . C. ciliatum, Dumontia filiformis, . Dilsea edulis, Furcellaria fastigiata, Lithophyllum lichenoides, . Corallina officinalis, C. squamata, C. rnbens, 5 | Feb., 191. | Roonah Pt. (Clare Tsland. | April, 1911, | July, 1909. Clare Island. Oct., 1910. Clare Island. x | | | | | | | ab. ab. | mS eS: OS eS 3% 3 SS Re SS Se Se 8 Se ab. ab. ab. os Od OS OS 26 OS 2h OK SDS OS OX OS BOOS x | | x x s mS OS OS Oh 8h RK SK KK BR Se ke Se aS PN OS OS OR OS ON he ON” Oke OM i} mS OS OS oo OM x x x xX X X xX 15 46 Proceedings of the Royal Trish Academy. 1. Lrposed Caves.—Several of these occur between Portlea and the light- house, and are accessible at low tide. The walls, almost up to the roof, are covered with encrusting organisms representing algae, lichens, and sponges ; and except for local patches, shrubby species are absent. In the following list, the first four are dominant, occurring roughly as bands from above downward in the order mentioned, whilst the three last are scattered irrecularly a short distance above high-water mark :— Hildenbrandtia prototypus. Lithothamnium laevigatum. Verrucaria sp. Ralfsia clavata. Lithothamnium Lenormandi. Cruoria pellita. L. polymorphum. Large sheets of sponges also occur, and amongst these Halichondria ponicea, frequently with Rhizoclonium Kerneri f. endozoica, is abundant. Small patches of a very short growth of the following fruticulose species are occasionally found, their presence probably indicating local shelter :— Phyllophora rubens. Plumaria elegans. Chylocladia ovalis. Corallina officinalis. Where fresh water exudes from the roof or walls, streaks of Cladophora rupestris, if the exposure be not too great, mark its presence, and at times Rhodochorton floridulum in addition. Pools are generally lacking; if present, their flora resembles that given below for sheltered caves. 2. Sheltered Caves.—Several types of these may be distinguished according to the nature of the shore and the manner in which they receive protection. In our area the two following only need be mentioned:—(a) Open caves on exposed coasts, protected by rocks at the entrance (Roonah Point) ; and (0) high-level caves with the entrance at, or just above, high-water mark. In the former there is often a variety of ground and vegetation, especially if boulders and pools occur; in the latter the conditions are more uniform. (a) The encrusting vegetation at Roonah Point is the same as that on exposed coasts, but less extensive; the mossy growth of Rhodochorton Rothii is abundant on the drier boulders and on the upper parts of the walls; Sphacelaria britannica and various Cyanophyceae occur with this species, and Phyllophora Traillii is found in patches. An abundant growth of shade- loving Florideae follows at a lower level, covering rocks, boulders, and shallow pools, Chylocladia articulata, Delesseria alata, Nitophyllum laceratum, Polysiphonia wreeolata, and encrusting species such as Lithothanniun Clare Island Survey— Marine Algae. 15 47 Lenormandi and L. polymorphum beng the most usual. The pool-flora consists chiefly of the following species :— Phyllophora rubens. Polysiphonia urceolata. P. membranifolia. Chylocladia articulata. Rhodymenia palmata. Griffithsia setacea. Nitophyllum laceratum. Pterosiphonia parasitica. Delesseria sanguinea. Lithophyllum incrustans. Plocamium coccineum. Lithothamnium polymorphum. (6) A small tunnel-like cave on the south side of Portlea was carefully investigated each season. Though short, it was the only really dark cave examined, and its flora proved of considerable interest, three of the rarest algae of the island occurring in it. Near the mouth of the cave the shaggy growth lining the walls consisted of Cladophora rupestris (dominant) and Plumaria (sub-dominant), with the following species scattered freely amongst them :— Ectocarpus Holmesii. Polysiphonia macrocarpa. Gelidium pusillum. Rhodochorton floridulum. Rhodymenia palmata. Callithamnion scopulorum. Polysiphonia urceolata. Spermothamnion Turneri. A little further in, Plumaria becomes dominant, and, with the exception of Cladophora rupestris, which assumes a thin feebly branched habit, the other algae gradually disappear. On the bottom and at the margins of pools, Derbesia marina is found, and on the sloping sides Codiwm amphibiwm under the drip of fresh water. At the further end, the walls are mostly clothed with a pure growth of Plumaria, though patches of Lithothamnium Lenormandi var. squamulosa exist, and Derbesia occupies the water-edges. The pool-flora is remarkable ; Plumaria, Ceramium acanthonotum and Polysiphoni” macrocarpa are common, whilst further in there is a felt of the minute Ptilothamnion lucifugum (sp. Nov.). (b), Sub-littoral Region. The sub-littoral vegetation of exposed coasts is difficult of exploration. In calm weather much may be learnt from a boat at dead low water, but in a general way a small boat and a hand-dredge are of little use. Two associations only are described below, these being the only ones which were definitely investigated. Colonies of the beautiful Cystoseira ericoides are plentiful off the shore, in company with such plants as Saccorhiza and Himanthalia, but unfavourable weather baftled attempts at their study. 15 48 Proceedings of the Royal Irish Academy. Laminaria Association. Clare Island is girdled at low-water mark with a fine belt of Laminaria, — which descends on all sides deep down into the sub-littoral region. With the exception of the somewhat doubtful Z. hieroglyphica, all the English species © occur, so that the west of Iveland presents the exposed Laminaria vegetation of the British Isles in its most varied form. Though a submerged association, each species reaches to the top of the sub-littoral region, and is thus visible during the lowest spring-tides. Alaria and Z. digitata are uppermost, and are often uncovered; Saccorhiza and ZL. saccharina follow, and usually show in calm weather, whilst the erect stipes of Z. Cloustoni protrude in favoured localities only. Alaria, Z. digitata, and L. Cloustoni form well-marked belts which are always to be found on the type of shore that they respectively require, whereas the growth of Saccorhiza, and ZL. saccharina, though at times plentiful, is scattered and local. A steep coast with almost perpendicular rocksis the A laria shore par excellence, though L. digitata is also usually present. On sloping rocks the same vegetation obtains, the amount of Z. digitata increasing as the shore becomes less steep, till at length Alaria disappears altogether. JZ. digitata often descends into many fathoms (at least 8); but Alaria appears to form a shallow belt in the uppermost part of the littoral region. Z. Cloustoni enjoys the Atlantic swell, but prefers a flat bottom, and is absent on steep rocks. Within the shelter of a reef, or in other similarly protected spots, the plants are found nearer the surface of the water, and at the lowest tides the tops of their stipes protrude. In such positions the association may be conveniently examined from a boat (e.g., at Roonah Quay). A flat or gently sloping bottom with a certain amount of shelter is most suitable for Saccorhiza and L. saccharvina. In certain spots on the south shore, and especially in channels between the rocks, the former grows in plot-like areas, which may be easily examined from the rocks in quiet weather. It is also plentful in the shallow water off Kinnacorra. Streak-like patches of L. succharina have also been noted in these localities, but the plant is more characteristic of quiet regions. As stated on p. 15, the Laminaria association, though it may be partially uncovered during spring-tides, marks in reality the upper limit of the sub- littoral region. The amount uncovered varies with the exposure. In the most boisterous spots Alaria ascends high, and consequently, if a spell of quiet weather synchronizes with spring-tides, a considerable amount, 5—5 feet, is left bare. Borgesen mentions a case in the Faerdes, where Alaria flourished at a level above that of high-water line. Nothing like this has been seen in Clare Island Survey— Marine Algae. 15 49 Ireland, and it probably only occurs where great exposure is coupled with a wet, foggy climate. The usual epiphytes characteristic of the species occur. Litosiphon Alariae, Ectocarpus fasciculatus, and Myrionemaceae are frequent on Alaria— Chantransia Alariae rare. The laminae of JZ. digitata commonly bear a copious growth of Ectocarpi, Ceramia, Callithamnion tetragonum, and other small plants, while the stipes are usually clean. The rugged stems of L. Cloustoni, on the other hand, are densely clothed with Florideae, which show a tendency to definite sequence from above downwards as pointed out by Berthold (82) and Bérgesen (05). As a rule this sequence is as follows :— Rhodymenia palmata. Polysiphonia urceolata. Delesseria sinuosa. Delesseria alata. Heterosiphonia coccinea. But in addition to the differences due to seasonal variations many irregulari- ties were noted. Other characteristic epiphytes are :— Codium tomentosum. Ptilothamnion pluma. Callophyllis laciniata. Ptilota plumosa. Rhodymenia palmetta. Plumaria elegans. Lomentaria articulata. Rhodochorton parasitica. Plocamium coccineum. Cruoriella Dubyi, r. Nitophyllum laceratum. Cruoria pellita 7. N. uncinatum (autumn). Petrocelis Hennedyi 7, N. Bonnemaisoni, +. Dermatolithon pustulatum, Saccorhiza is usually fairly clean, as is also ZL. saccharina in exposed localities. Comparison with other Countries.—As far as Alaria, LZ. digitata, and L. Cloustoni are concerned, the above vegetation agrees well with that described by Borgesen for the Faerdes, though in that region it ascends somewhat higher on the shore. Saccorhiza is entirely absent; but this is compensated for by the presence of two additional species, A. Pylait and L. faeroensis, the latter being confined to sheltered coasts. Bérgesen, in an interesting paragraph on the effect of wave-action on Laminariae, points out that the pliable stipes and fronds of Alaria and JL. diyitutu are able to resist the violent conditions of a steep shore, and where the breaking of the waves is felt; whilst the stout and comparatively rigid stems of LZ. Cloustoni are suited to the strong pull of the rollers in deeper water. R.I,A, PROC., VOL, XXXI. G 15 15 50 Proceedings of the Royal Irish Academy. _ The Swedish vegetation described by Kylin is very different. Alaria is absent, and the verticai rocks are clothed with Z. digitata and L. saccharina var. bullata. In the most open spots LZ. digitata var. cuneata is dominant. The Z. Cloustoni vegetation is entirely separate from Z. digitata, being found only in the deeper part of the sub-littoral region, appearing first in about 15 m. of water. Boye’s account of the Norwegian flora (according to Boérgesen’s summary) agrees with that of the Faer6es; and Kylin’s notes on the vegetation near Bergen (10) indicate the preponderance of Alaria and L. Cloustont. The Iceland vegetation resembles that of the Faerées, but is enriched by the presence of L. nigripes and Saccorhiza dermatodea. Tn the south of England Alaria is rare except in Cornwall, but apart from its absence the Clare Island vegetation agrees well with that noted in Dorset. The exposed rocky coasts of Ireland are, however, more favourable for an extensive Laminaria vegetation, and are highly suitable for a growth of Alaria, Encrusting Alga Vegetation (sub-littoral). The Lithothamnium vegetation of the littoral region extends downwards into the deepest parts of the sub-littoral, but, owing to the difficulty of dredging amongst a dense growth of Laminaria digitata and L. Cloustoni, it is difficult to ascertain the amount of growth and the relative abundance of the species. An extensive coating of calcareous algae is, however, present, and on exposed coasts probably occupies a large part of the available rock-surface, though soft encrusting species, such as Cruoria pellita, Cruoriella, and Peyssonnelia, occur with it. Lithophyllum incrustans enters the sub-littoral ; but, as far as could be seen, the Lithothamnium ‘band at the lowest tide-level consists mainly of L. polymorphum, and this extends downwards for several fathoms. Other species dredged were LZ. laevigutum and L. compactum. The fruticulose calcareous algae, such as L. calcareum and L, fasciculatwm, are characteristic of sheltered shores (p. 69). B.—Sheltered Coast Series. (a) Littoral region. Several of the associations found on sheltered coasts have been already dealt with. In some cases the principal differences noticeable in passing from the open shores to quieter conditions have been pointed out, so that re-descrip- tion is unnecessary. In other cases, though’ theassociations have been described, their growth in sheltered localities requires special notice; yet other communities are peculiar to such regions, Clare Island Surveyj—Marine Algae. 15 51 Hildenbrandtia—Verrucaria Association. Brief notes may be added to previous remarks with regard to this association (p. 19). On the sheltered shores of Clew Bay the same species are present, especially on boulders and smooth rocks. Where these are absent a discontinuous growth on stones and pebbles occurs, and in such spots Hildenbrandtia is joined near low-water mark by Porphyrodiscus simulans and Fihododermis elegans (plants hardly distinguishable in the field), and also by Ralfsia clavata. H. prototypus itself, moreover, appears to extend to the sub-littoral region (cf. p. 20). The difficulties of obtaining satisfactory material and of distinguishing the species when collecting, make it no easy matter to gain a true idea of the different societies. It is certain, however, that in these localities the Ralfsia clavata vegetation is one with the Hildenbrandtia association, and that it forms a part of it. It is further evident that a vegetation composed of soft encrusting algae and lichens extends uninterruptedly from high-water mark down to the Lithoderma association of 5-8 fathoms. The details require working out, but roughly the principal species form zones as follows :— Hildenbrandtia prototypus (upper littoral). Verrucaria mucosa (middle littoral). Ralfsia clavata (lower littoral). Cruoriopsis Dubyi (shallow sub-littoral). Lithoderma fatiscens ? (shallow sub-littoral). Fucaceae Association (sheltered). The growth of Fucaceae on flat sheltered shores is exceedingly luxuriant, and often forms a conspicuous feature of the landscape. The indispensable factor is good anchorage. On flat shores, where wave-action is slight stones are sufficient to supply this necessity, but where rocks and stones are wanting Fuci are conspicuous by their absence. The association was studied especially in Achill Sound, the north part towards Inishbiggle and Bellacragher Bay having been examined, as well as the southern portion towards Achillbeg. As far as was seen the Fucus growth in Clew Bay is of a precisely similar nature. The association is composed of the usual five species in their normal sequence (see p, 23), and where fresh-water streams occur, the sixth species, F. ceranoides, invariably appears. In some places the zones are fairly sharply defined ; in others they are much less distinct, and each zone overlaps the next. A certain amount of irregularity exists too as to the order in-which Ascophyllum and 2. vesiculosus occur, The former usually commences above G2 16 52 Proceedings of the Royal Irish Academy. the latter; but not infrequently /. vesiewlosus forms a very wide zone reaching from F. spiralis down to F. serratus, with Ascophyllum scattered amongst it, or forming more or less of a band in the upper part. The reason for this is explained later. At times one or more of the species may be absent, and this may almost always be traced to lack of suitable means of attachment at the appropriate level. In Achill Sound, for instance, the association often stops abruptly with Ascophyllum or F”. vesiculosus, owing to the absence of stones at the level at which F! serratus should occur. In other places Pelvetia and I’ spiralis are only feebly developed owing to the lack, in the upper part of the shore, of stones of sufficient size. Some idea of the width of Fucus association in Achill Sound will be gained from the following readings :— A. Sloping shore near Achill Bridge with a narrow band of Fucaceae :— Width of zone. Pelvetia, . ; ; ; F ‘ ; . 2-8 feet. F. spiralis, ‘ : 5 : : : ae ND=6 Be F. vesiculosus, . ‘ : ; . é . 10 a F. vesiculosus and Ascophyllum mixed, ‘ a al) 3 F. vesiculosus, . j j 7 ‘ : ere al) FA F. serratus, ‘ j ; ; ; ; « 210 Pr B. Flat shore with a very wide Fucus association ; about 1 mile south of Achill Bridge :— Width of zone. Pelvetia, i : : 6 j n : 5 yards. EF. spiralis, : ; ; ; ‘ ; > LOM IRS Ascophyllum, . ; : ; : : « 64 OCS Ascophyllum and F’. vesiculosus, ; : at (OOR ES F. vesiculosus, : 6 c : ; s, Ole F. vesiculosus, F. serratus, . F : . oO) ep The above figures represent fair average readings for sloping and flat shores respectively, and serve to show that the horizontal range varies directly with the amount of slope. On very flat shores even wider bands exist. At Mulranny, for example, on a flat stretch of rocks and stones at high-water level, a growth of Pelvetia 80 yards wide was noted: this was followed by a sudden increase in the fall, and the succeeding band of & spiralis was only 3-4 yards in width. With regard to exposure, all the species composing the association can tolerate the extreme shelter of the inner part of Clew Bay. On passing from this region to the more open, beyond a slight change in form, there is no Clare Island Survey—Marine Algue. 15 53 inarked difference in the vegetation as long as Ascophyllum remains. A slight exposure, however, is sufficient to affect this species, and with its disappearance the association is robbed of one of its most conspicuous elements. Ascophyllum begins to disappear on the outer sides of the islands, but persists in all cases on the inner sides. West of the islands it is only found in certain places, though this perhaps is largely due to the lack of good mooring. But, in spite of this, there is no question whatever that in the Survey-area Ascophyllum is far more susceptible to exposure than F. vesiculosus. This is in agreement with observations in other parts of the British Isles, and also with accounts that have been given of the Norway Fucus-vegetation (Boye, 94, and Kylin, ’10). The conditions described for the Faerces, where Ascophyllum is the less susceptible of the two, are very remarkable, and, as far as known, unique. The result of exposure in producing a change in form in the various constituents of the association is very marked. It is best seen in F. spiralis. F. vesiculosus also produces a wealth of form, but the variations are more difficult to connect with external conditions. Remarks on these points occur in the notes on each species given later. Fucus Farms.—The botanical features of the plots, planted with stones in order to obtain a growth of Fucus for use as manure, may be dealt with here. The farms are formed on flat stretches of sand exposed at low water. They may either extend from the shore, or be laid out as isolated patches in the middle of the strand. The rapid colonization, and the luxuriance of the subsequent vegetation, prove beyond question the effect of anchorage. The stones are covered mostly by £. vesiculosus, though a small amount of Ascophyllum sometimes occurs. At the lowest levels there are occasionally encircling belts of F. serratus, but the stones are not as a rule placed sufficiently low for this species. Ff. vesiculosusis very luxuriant, forming large tufts, bright yellow in summer and deep green in winter, with numerous bladders and abundant fruit. 2 serratus, when present, grows equally vigorously, with fine broad fronds often a yard in length. The plants are cut in February after two full years’ growth, and the stones turned over for a new crop of sporelings to develop. It is somewhat remarkable that this artificially produced vegetation con- sists of a practically pure association of F. vesicwlosus, forming thus a striking contrast to vegetation obtaining on mud-flats (eg., Belfast Lough, South-. ampton Water). The latter are colonized by mussels, and these support a pure growth of Ulva Lactuca. Ulva, and many other adaptable algae, are present in Clew Bay, but Fucus has complete sway on the stone plots. It would be interesting to know whether Fucus would develop if stones of sufficiently large size could be maintained on the mud-flats of Belfast Lough. 15 54 Proceedings of the Royal Irish Academy. A few notes are appended on the species composing the association. (a) Pelvetia canaliculata.—This is less variable than any other species. In the sheltered parts of Clew Bay and in Achill Sound, it usually occurs on the irregularly scattered stones, and consequently the zone is interrupted. The growth is entirely flooded at spring-tides, but at neap-tides the upper part of the belt is unmoistened, the plants remaining quite dry, and brittle for several days in succession. Though Pelvetia may form a zone several yards wide, its vertical distribution is not more than 14-2 feet. It fruits freely during summer and autumn; but the growth being more interrupted and not so Inxuriant as on open shores, the orange-yellow band is not so conspicuous. (b) F. spiralis—Not only is F. spiralis found in all the sheltered parts of Clew Bay, but it is capable of growing on peat and loam, and ranges from such situations to the roughest and most exposed parts of the open coast (cf. Borgesen, ’05, p. 746). Leaving aside its growth on peat (see p. 124), we find that on flat shores the plants are usually stone-attached, and do not form so dense a vegetation as when growing on rock. For this reason they become drier during the long exposure to the air, and are shorter and less branched. When the shore is composed of rocks and large stones, the growth is more vigorous; and under these conditions the plants attain their maximum size, the fronds being wide, much branched, and bearing large fleshy conceptacles. This form remains as long as the conditions are sheltered, but when exposure is experienced, or where a current strikes the shore, the plants become shorter, though they usually retain their bushy habit. The belt formed is much wider than on steep open coasts, and its width is proportional to the degree of slope. In Achill Sound it varies from a few feet to 10 yards, and usually forms a pure growth, though /. vesiculosus and Ascophyllum occasionally intrude into the lower part. The fruiting season on quiet shores appears to extend over a slightly longer period than on those which are more open. In Clew Bay it commences in May, and does not cease till autumn. On sheltered shores also, a difference in colour is noticeable during the autumn, the plants turning a bright rusty brown. The same peculiarity has been observed on the south coast of England; and it serves to distinguish at a glance F. spiralis from F. vesiculosus, the latter being usually yellow or only slightly brown. In some places two sharp zones of colour are produced. Another feature, which on flat shores is very characteristic of F. spiralis, is the presence of long blister-like swellings on the fronds. ‘These are invariably present in the inner part of Clew Bay, and are entirely absent from the short forms found on Clare Island. Sauvageau has lately examined I, spiralis (= F. platycarpus, Vhur.) with great thoroughness (’08). Clare Island Survey—Marine Algae. 15 55 (ec) Aseophyllum and F. vesiculosus—The irregularities in the vertical range of Ascophyllum and F. vesiculosus have been noted by several writers. Borgesen calls attention to the difference in the observations of Strémfelt in Iceland, and Rosenvinge in Norway; and records irregularities noted by himself in a single locality in the Faerdes. He comes to the conclusion that the difficulty may be explained by the factor of exposure and small differences in local topography. No doubt this solution is correct with regard to many localities; but in Achill Sound it was found inadequate, and a further explanation had to be sought for. It has always been assumed that in England the Ascophyllum zone begins slightly above that of F. vesiculosus, and Miss Baker (’09) gives figures which support that view. This sequence is found on the stone embankments of Achill Bridge and in other places; but on the shores in Clew Bay, as often as not, the reverse order obtains. The explanation, so far as the sheltered waters of the Survey area are concerned, was found to be largely one of anchorage. Ascophyllum requires a stronger mooring than Fucus; and on the sandy shores of Clew Bay where the stones in the upper part of the beach are small they are insufficient for Ascophyllum, though well able to provide a mooring for a short growth of Fucus. The absence therefore of the Ascophyllum-zone, or the presence of scattered plants only, is explained by the absence of rock and large stones at the right level, or the presence of isolated rocks only. In many parts of the Sound the anchorage is sufficiently good to allow Ascophyllum to flourish at all levels, and then the normal sequence is almost invariably maintained. Very small stones suffice for both F. spiralis and F. vesiculosus; but it is unusual to find even young plants of Ascophyllum on stones. The last-named shows a decided preference for boulders, from which it hangs in festoons. The stone-attached Fucus-vegetation is essentially one of very sheltered water, and could not exist were there much wave-action. Where the stones are light the growth is small, and if, by reason of a long spell of quiet weather, an extra amount of foliage should be produced, a number of plants will be displaced with the first ruffling of the water. Ascophyllum fruits in late winter and spring. In summer and autumn a large amount of vegetative growth takes place; and young receptacles are laid down in autumn. The latter begin to ripen in March, the main crop of spores being liberated in April and May. The effect of exposure has been dealt with above. ; The Fucus vesiculosus belt is the widest. On flat shores it ranges from 20 to 80 yards, in the upper part usually accompanied by Ascophyllum, but near low water often pure. No attempt was made to determine all the forms ; but the most abundant is the ordinary type. Var. vadorwm is also common, 15 56 Proceedings of the Royal Irish Academy. and var. Jaterifructus not infrequent. In many localities var. a@illaris is found, which under certain conditions passes into F. volubilis, Huds. The latter in its typical form was not noted in the area; but the salt-marsh plant var. baltica is abundant. This, together with a peculiar variety named muscoides, is dealt with later (p. 80). (d) Fucus serratus. Whilst Ascophyllum enjoys hanging from boulders, F, serratus prefers to lie flat. Occasionally it is found growing on sand-covered rocks, and then rests on the bare sand. It is found on all sheltered coasts where there is appropriate anchorage; but in many parts of Clew Bay this is wanting; and the pebble-attached association sets in below F. vesiculosus, F, serratus forms a band which unites the littoral and sub-littoral regions. The fronds are wider on sheltered than on exposed coasts ; and on the whole it is the least variable Fucus. Sub-vegetation— When growing on sloping rocks a luxuriant undergrowth of various algae is found; but when attached to stones, or where much sand occurs, extraneous species are scarce, Amongst those almost constantly present are :— In upper part. In lower part. Hildenbrandtia. Cladostephus spongiosus. Catenella. Ulva. Gelidium pusillum. Ceramium rubrum. Cladophora rupestris. Chondrus. Gigartina. C. rupestris, On the more open shores where clean rocks are prevalent, the Laurencia- Gigartina vegetation often forms the main undergrowth, though upward extensions of the Corallina-Lithothamnium association are also frequent. Amongst these,such sub-littoral or shade-loving plants as Chylocladia articulata, Delesseria alata, and Plumaria occur, with many odd plants of various communities. In sandy places a Rhodochorton floridulum vegetation is often developed, and this ousts many of the other species. Epiphytes—These are likewise fewer in very sheltered than in moderately open localities. In the quiet reaches of Clew Bay, Pylaiella is always present, and forms in spring and summer large floating masses. Other frequent epiphytes are :— Dictyosiphon foeniculaceus. Ectocarpus siliculosus. Ceramium rubrum. EK. tomentosus. C. Boergesenii. E. fasciculatus. In late summer blue-green algae are much in evidence, Calothrix confervicola and Ltivularia atra occurring everywhere, often accompanied by Clare Island Survey—Marine Algae. 15 57 large masses of Lyngbya majuscula. The more open parts of the bay show, with the exception of Dictyosiphon and Cyanophyceae, which are rare or absent, the same epiphytes (see also p. 25). For the comparison of the present association with that of other countries, see p. 26. The Lichina Association (sheltered). Locally distributed on sheltered and semi-sheltered localities (Clew Bay, Achill Sound, Bellacragher Bay). See notes on p. 26. Porphyra Association (sheltered coasts). The growth of Porphyra found in many sheltered places, though not noted by previous writers, is very characteristic and worthy of record. It is found on two types of shore, viz. (1) gently sloping beaches of low rocks and stones, and (2) sandy bays where half-buried rocks are present. The growth occurs more or less throughout the year, though it appears to be more abundant in spring and autumn than in summer. On both shores it is usually accompanied by various forms of Hnteromorpha intestinalis and LE. linza. On beaches composed of stones and flat rocks the association usually fills up the gaps between Fucus. The latter is confined to the larger rocks, whilst Porphyra, which extends from below /. spiralis down to low-water mark, occupies the smaller stones and rocks, though it also occurs epiphytically on Fucus. The amount of growth noted varied considerably on different occasions, as did also the size and form of the plants. In summer it is mostly confined to the lower levels, where it may form a distinct belt. This type of growth is well seen at Louisburgh (immediately east of the Bunowen River), also on the inner side of Bartraw and on Annagh Island, where wave-action is entirely absent. Other parts of Clew Bay, which appeared equally suitable, were entirely devoid of Porphyra. [n the localities named a strong current runs along the shore, so that the conditions though quiet are not by any means stagnant, and this fact may in part explain the vigorous growth, and its absence elsewhere. The second type of growth referred to, namely that on rocks in sandy bays, is somewhat remarkable, because the expanded membranaceous thalli have to endure the constant friction of the sand. In spite of this, however, a very vigorous and characteristic vegetation is found. The growth is most notice- able at the sides of low bare rocks or those crowned with a mass of Fucus. It may be seen thus in many spots in our area, and has been noticed in other parts of the British Isles. But in addition to this, Porphyra occurs on rocks more or less hidden by sand, so that at first sight the growth might be taken for loose fronds that have become embedded. The plants are in reality RIA. PROC., VOL. XXXI, H 15 15 58 Proceedings of the Royal Irish Academy. attached to low-lying rocks, and are thus not only exposed to constant sand- movement, but are liable to be completely buried. On the other hand, with certain winds, nearly all the sand is removed by the tide; and the thalli lie on the bare rock. They are also liable to be covered by blown sand. Under average conditions it would appear that only a few inches at the base of the fronds are buried, and that the upper part remains free, lying flat on the damp ground: though one must assume that germination took place on rock. Associated with Porphyra are Enteromorpha spp., especially the large flat forms approaching £#. linza. This plant oceurs in the damper positions, being most abundant in the lower part of the association, and reaching down to the sublittoral region. In August the Enteromorpha growth was in all cases much less than in May. The zone formed by Porphyra on sheltered coasts is more extensive than in the open. In the sandy bays described above the plant escapes a certain amount of desiccation, and the result of this is seen in the fact that whilst on bare rocks it does not exceed the /. spiralis level (at all events during summer), on sand-covered rocks it frequently reaches Pelvetia. ‘lhe lower limit also is further than that of the exposed association, for the plant is frequent at low-water mark, and occasionally extends well into the sub-littoral region. The form assumed in sheltered localities is that.of the divided and lanceo- late type, var. daciniata J. Ag. his is specially well developed on damp rocks and on margins of pools. On drier rocks and with increased exposure there is a tendency for the fronds to become short and umbilicate. The sand- rock form of the association is well seen at Mulranny, also between Old Head and Leckanvy, whilst a small patch exists in the bay on Clare Island. Though not noted, forma linearis probably exists in winter in the upper levels, as in exposed localities. A similar vegetation of Porphyra would, judging by the localities described by Rosenvinge, appear to be widespread in Denmark (’09, p. 65). Laurencia-Gigartina Association. Fairly well developed in many parts of Clew Bay outside the islands. See p. 39. Corallina-Lithothamnium Association. At times feebly developed in the more open parts of Clew Bay. See Be), _ Corallina-Cladostephus Community. In company with the Laurencia community of sheltered coasts there often occurs a similar vegetation of short dense growth, but composed largely of Clare Island Survey—Marine Algae. 15 59 Corallina, Gelidium, and Cladostephus, and having a tendency to approach the Rhodochorton floridulum association. The presence of Cladostephus spongiosus together with Corallina officinalis is very characteristic of this vegetation, and hence the name selected. The community in question is essentially one of flat, sheltered rocks, avoiding the clean rocks and turbulent conditions enjoyed by Laurencia, but thriving with more slope and slightly more exposure than R. floridulum. It occupies the same bathymetric range as Lawrencia pinnatifida, and though capable of growing in the shallow sub-littoral region, it more often gives place to Gigartina or Laminaria. At Portlea it is intimately associated with Rhodochorton, but the two communities are usually sharply defined, and distinguishable at once to the eye, as well as to the tread. Though both Corallina and Cladostephus are rock-attached, a thin layer of sand is often found on the surface of the substratum. Corallina, under the conditions obtaining, is short and more or less erect, its stiff branches easily retaining sand and fine debris, so much so that the presence of blackened sand and particles of decayed matter is almost characteristic of the community. At the same time it is not truly psammophilous. A vegetation of the same nature (though differing slightly floristically) has already been described by me in connexion with the growth of Ulva, and the origin of the blackened sand is explained (11, pp. 124 and 128). The Corallina-Cladostephus sward is seldom so extensive as that of Rhodochorton floridulum, being patchy, and only occasionally forming broad belts. Gelidium aculeatum is sometimes present in quantity, and C. sqwamata is common, especially in the more open regions. A very heavy epiphytic vegetation exists, ef which Hnteromorpha elathrata, Ulva, Leathesia, Coramium flabelligerum, and C. rubrum are the most abundant. Callithamnion Hookervi also is found in summer on Cladostephus. In addition to the above, a large number of sporelings occur. These succeed better on this vegetation than on Rhodochorton (where sporelings are exceedingly numerous); and though the plants do not attain their maximum size, they usually fruit, and are sufficiently large to give distinct patches of colour. Much Cladophora sericea and C. albida is present in spring ; Ceramium rubrum in various forms is abundant in summer; whilst in October (1910) a form of Hnteromorpha clathrate was very conspicuous. In summer Ulva also is present in great quantity, as has already been pointed out (Cotton *11, p. 128), being saxicolous or epiphytic. On Clare Island the association occurs on a small scale at Portlea and Kinnacorra; and on the mainland at many spots in the outer part of Clew Bay (Roonah, Louisburgh, Mulranny). A similar vegetation occurs on the H2 15 60 Proceedings of the Royal Irish Academy. south coast of England; and though not recorded from other countries, it probably exists, at all events, on the north of France. Rhodochorton floridulum Association. Small patches of this are occasionally found on rocky shores; but it is characteristic of the sand and low-rock series described later (p. 65). Sand-Pool Vegetation. Occasionally met with, but typically developed on sandy shores (see p. 67). Cave Vegetation. The caves of sheltered coasts are dealt with on p. 46. (b) Sub-littoral Region. Laminaria Association (sheltered). The Laminaria vegetation of sheltered shores is fairly well developed. It is found in many parts of Clew Bay, in Bellacragher Bay, and in parts of Achill Sound. The characteristic species are L. saccharina and L. digitata, the latter being, as a rule, confined to localities where current is present. In the deeper and more open parts, Z. Cloustont exists; but with the appearance of this species Z. saccharina drops out, and the association passes gradually into the exposed type. Whilst on exposed coasts the Laminaria vegetation is typically in belts, on sheltered coasts it is commonly discontinuous, and occurs rather in plots. This is mainly owing to the scattered nature of the anchorage. It is found wherever rocks or large stones occur; and the amount is determined by the extent of the stony area. In Clew Bay, the hard bottom is found in three kinds of ground, each of which supports a different type of Laminaria vegetation. As the latter are determined by definite ecological conditions, and as, in Clew Bay, they scarcely overlap, it is reasonable to regard them as three sub-associations. The sub-association of Laminaria saccharina (most often var. latissima) oceurs in the inner part of the bay, where the conditions are quiet and the current weak. The fronds are long and very broad—in extreme cases up to a yard across. Occasionally the association is pure (except for small species forming the undergrowth); but as a rule other algae intrude, such as Cystoseira granulata and Halidrys siliquosa. Rarely the association merges into the Zostera beds. The second sub-association is characteristic of ground Clare Island Survey—Marine Algae 15 61 where there is a strong current, and is found typically in the channels of 3 to 5 fathoms between the outer islands. It consists entirely of L. digituta. The bottom is scoured by the tide; but the larger stones and blocks of rock support a vigorous growth. The plants are remarkably strong, and in the deeper parts the fronds often attain a great length. Epiphytic species are few ; but in such localities the peculiar plant known as Rhodymenia palmata var. sarniensis is occasionally met with. The third type of Laminaria vegetation, characterized by the presence of ZL. Cloustoni in addition to L. digitata, is found only outside the islands. It extends over large areas in about 7 fathoms, and passes gradually into the exposed vegetation that occurs further out. Representatives of these same sub-associations are found elsewhere. L. digitata is abundant near the south entrance of Achill Sound ; andin the quiet waters of Bellacragher Bay a distinct ZL. saccharina vegetation exists, which in several places extends to the shore. In other counties, the sheltered type of Laminaria vegetation has not received much attention. In the Faerdes it is well developed, and shows similar variations to those in Clew Bay. though with floristic differences. Z. jfacroensis takes the place of L. saccharvina in quiet bays, and L. Cloustoni forms the vegetation of channels. The Laminaria zone described by Joubin (09) includes patches of LZ. saccharina attached to stones ou a soft bottom, which obviously correspond to the sub-association of that species dealt with above. Encrusting Alga Vegetation. Occasionally found, but more often on exposed coasts (see p. 50). Il.—Sand and Sandy-mud Formation. The present formation is characterized by a large quantity of movable sand or sandy-mud, in which the vegetation is liable to be partially or completely embedded. In its extreme form, the shore consists entirely of this matrix, the algae being immersed in sand, forming, in fact, a sand-binding community. But the sand formation passes by transitions into the rock-formation on the one side, and to mud-flats on the other. As an example of a-sand-binder we may take Microcoleus, whilst Vaucheria Thuretti supplies an instance of a plant that fixes sandy-mud. Both form extensive associations. Rhizoclonium riparium also may under certain conditions form a surface-covering on a shore of pure sand. In the majority of sand-beaches, however, all vegetation is absent and only begins to show when solid ground is reached. Where low half-buried rocks occur a 15 62 Proceedings of the Royal Trish Academy. psammophilous vegetation is found, of which Rhodochorton is a good example. With extreme shelter, this can form a mossy carpet on pure sand; but as a rule it is only found on sand-covered rocks, or on small intervening sand- spaces. The alga is probably in the first instance attached to rock, but as the plants grow they form adventitious rhizoids and accumulate sand. Being saxicolous, Rhodochorton extends to rocky shores, where sand is scarce or even absent. As an association, however, it is only found where this material is plentiful. Several other psammophilous species accompany Rhodochorton. The Corallina-Cladostephus community represents the last stage m the transition from sand to rock. Though frequent on sandy shores, a bed of rock is essential for its growth, hence it has been dealt with under the rocky- shore formation. The sand-pool vegetation is, on the other hand, described in the present section, as the characteristic species, though attached to rock, are markedly psammophilous. The sub-littoral associations call for no special comment, except that the last three are not strictly psammophilous, though in our district they are confined to sandy areas. They are placed under this heading for convenience, and therefore provisionally. The associations and communities recognized in the sand and sandy-mud formation are as follows :— (a) Littoral Region. Rhizoclonium ripariun. Microcoleus chthonoplastes. Vaucheria Thureti. Vegetation of vertical banks. Rhodochorton floridulum, Corallina-Cladostephus. Porphyra umbilicalis. Sand and sandy-mud series : | Sand and low-rock series : ! Sand-pool vegetation. (b) Sub-littoral Region. Rhodochorton floridulum. Zostera marina. Fruticulose Lithothamnium. Pebble-attached association. Encrusting algae. Channel-vegetation. Clare Island Survey— Marine Algae. 15 63 (a) Littoral Region. Rhizoclonium riparium community. The only community found on pure sand in our area is that of Rhizoclonium, though for this also a small amount of binding material is advantageous. The vegetation is found immediately below high-water line, and is occasionally very conspicuons, specially in spring and summer. At Mulranny, where the shore is unusually flat, Rhizoclonium covers extensive areas, patches an acre in extent having been noted.! The underlying sand (or sandy-mud) is for the most part firm, often bound together by Microcoleus, but in other spots it is loose and apt to become dry. In either case Rhizoclonium lies as a loose covering, thus feeling the effect of wind and tide, often being more or less buried. In some localities the upper part of the association is during neap-tides not wetted at all, and in this way differs from the somewhat similar filamentous vegetation which les on wet mud, e.g. on the Vaucheria Thureti association in Achill Sound (p. 64). Rhizoclonium ripariwm is usually a pure dominant, but amongst it (especially in the damper parts?) occur Hnteromorpha prolifera; E. clathrata var., and Cladophora fracta. The blue-green species, so common in the corresponding association on the salt-marsh, are absent. A very similar type of growth to that described above has been noted in Poole Harbour, and on a small scale near the mouth of the River Bann. (See also salt-marsh formation, p. 79.) Microcoleus chthonoplastes association. Of the sand- and mud-binding algae one of the most important is M/iero- coleus chthonoplastes, a microscopic species of world-wide distribution. The plants are often completely hidden in the substratum; but their presence can be detected by the way in which the soil holds together, and peels off in crusts, when lifted up. Though Microcoleus has not been previously recorded as forming an association as such, the plant-community has as a fact been long known, and for a description one cannot do better than quote Gomont’s note in his revision of the Oscillarieae (09, p. 92) :— “Le Microcoleus chthonoplastes se rencontre quelquefois a V’état de filaments isolés mélangés 4 d’autres Algues, ... mais il occupe aussi a lui seul de vastes étendues sur les cétes marécageuses ou sablonneuses. II] joue un role des plus utiles dans l’exploitation des marais salants en recouvrant le fond des bassins ou ocillets V@un tapis compact qui permet d’enlever les cristaux 1 The flatness of the shore at this spot may be judged from the fact that the Pelvetia zone, which is only about 1 foot deep, is in places oyer 80 yards wide. 15 64 Proceedings of the Royal Irish Academy. de sel sans mélange de vase. En cet état il forme des assises diversement colorées, accumulées sur une épaisseur parfois considérable et dont les supérieures, qui sont les seules vivantes, peuvent étre détachées par larges plaques et roulées comme des morceaux de drap. Ce végétal meérite ‘done complttement le nom de chthonoplastes (constructeur du sol) qui lui a été donné par les premiers observateurs.” In Clew Bay the Microcoleus association was usually found where there was a certain amount of mud. It occurs in large patches at Mulranny and also in the Belclare neighbourhood. In the small bay to the north-west of Achill Bridge it forms a broad band just below high-water line, binding the upper layer of mud into a firm sheet. It was also noted in several localities in fairly pure sand, but only as covering small areas. Various other blue-green algae enter into the composition of the association, notably :— In upper parts : In lower parts : Lynebya aestuarii. Vaucheria Thuretii. L. lutea. Rhizoclonium riparium. Microcoleus tenuis. Enteromorpha torta. Vaucheria Thuretii Association. This vegetation takes us further still from the pure sand, being found only where there is a considerable proportion of mud. Future work will probably show that it should be regarded as an association in the mud-formation. V. Thuretii forms a dense mossy growth embedded in the matrix, and appears to cover very extensive areas. Several samples were taken both from Clew Bay and from the muddy inlets north of the bridge at Achill Sound; and in each case the mossy tuft consisted of this species, often in beautiful fruiting condition, and practically pure. The growth is always immersed, the tips of the filaments alone showing, thus differing markedly from V. coronata, which forms a clean velvety growth. Felts composed of Rhizoclonium, Chaetomorpha and Cladophora fracta often lie on top of the Vaucheria, and these, in many cases, appear to be attached, and not merely drifted fragments; so that it is possible that careful study would reveal definite societies of other species amongst the general vegetation of Vaucheria. : The vertical range of V. Thuwretii is determined by the extent of the sandy mud. It flourishes in the lower part of the littoral region, and probably descends to low-water mark. Its upper limit is usually immediately below F. spiralis, but whether this was due to the lack of sufficient mud or to other causes in addition (such as long exposure to the air) was not ascertained, On 4 Clare Island Survey—Marine Algae. 15 65 the banks of rivers and where the water is brackish a zone of V. coronata is the characteristic vegetation near high-water line, but in Achill Sound that species is absent, and Microcoleus chthonoplastes is the dominant alga. Vegetation of Vertical Banks. Vertical peat-banks, abutting on the shores of Clew Bay and Achill, exhibit a similar vegetation to that described later for the salt-marsh formation (see p. 82). Rhodochorton floridulum Association. This association, though not hitherto described, is very distinct and of common occurrence in many parts of the British Isles, including the Clare Island district. tis characteristic of sandy shores where rocks are present, and is found on moderately open as well as sheltered coasts. Lhodochorton joridulum is the most important of a group of finely branched, upright-growing algae, which retain quantities of sand, if they do not actually bind it together, Other members of the group are :— Cladophora arcta, Ceramium strictum, Polysiphonia fibrata, C. cilatum. In suitable localities the association extends from the Fucus spiralis belt down to low-water line, but it also descends into the sub-littoral region, being found abundantly between the islands in Clew Bay. It prefers flat or slightly sloping shores; on steep shores the sand is washed away, and the plant is practically absent. Being of small size, Rhodochorton cannot fix sand on the same scale as phanerogamic plants, nor does it occur on sand apart from rocks! except in very sheltered areas, or where it issubmerged and beyond the effect of rough water. It is, however, capable of covering flat or sloping rocks with a dense mossy growth; and of carpeting the intervening sand-spaces so that they completely resist normal wave-action. In its young state the plant is probably attached to rock, but as growth proceeds, sand collects, and the abundant production of rhizoidal filaments helps to retain it. The growth forms either pads which may measure a foot or so across, or vast sheets more or less interrupted, but extending at times over a very wide area. When well- grown, the Khodochorton sward is about 3cm. high; but in favoured spots pads up to 6cm. have been met with. Though sometimes exposed to the full light, especially on the lower part of the shore, Rhodochorton grows best with partial shade. But even when exposed to light it is the ends of the branches only that feel the full effect of sun and air, as the base of the plant is covered with sand. B,I.A, PROC., VOL. XXXI. I 15 15 66 Proceedings of the Royal Irish Academy. Of the other species entering into the association Polysiphonia jibrata, Ceramium strictum, C. ciliatwm form conspicuous patches in spring and summer, their tufts of rhizoids growing down into the sward, and attaching themselves to the sand-grains. @. ciliatuwm is found more or less all the year round, and is of a shorter and denser habit than when growing in pools. In spring the © association often assumes a bright green colour due to the presence of Cladophora arcta, This species retains the sand well by means of hooked branches and rhizoids which grow downwards from the main shoots. It occurs not only within the Rhodochorton vegetation, but outside it, often forming on its flanks large unmixed patches. A dwarf form of Sphacelaria cirrhosa is also frequent, but, because of its small size, is apt to be overlooked. Though a filamentous species, it is distinct from the others, being attached by a basal disk, and not by rhizoids. In late summer, especially in the more sheltered localities, Rhodochorton is much discoloured by the presence of cyanophyceous epiphytes. Of these Calothrix confervicola is the most common, being at times so abundant as to ereate large black patches. Aphanotheca pallida is also frequent. On the margins, where the association joins the Corallina- Cladostephus vegetation, many other algae appear, and it is probable that the borders of the two communities are in a continual state of change. Young plants of Ulva Lactuca are at times present in great quantity, especially in spring, but this species is in no way a sand-binder, possessing only a foliose thallus and a basal disk. In the same way we may dismiss many other intruders, the Rhodochorton sward being, in fact, a general nursery ground, the sporelings succeeding each other in regular sequence as the seasons come round. With regard to distribution, the present community is probably very widely spread in the British Isles. hodochorton floridulum is one of the few species that form extensive associations on the flat sandy shores of Sussex; and it has been found growing in the same way in Dorset and Devon. Harvey states that at Kilkee (Co, Clare) “it often carpets large spaces on the rocks” ; and Ralfs writes in a similar strain as to the plant at Land’s End (in /itt.). It is also abundant on the shores of Belfast Lough. With regard to the Survey area, the association is found at Portlea on Clare Island (rocks giving the necessary shelter), and on a small scale in several other bays, where protection is afforded by a heavy growth of Fucus. On the mainland, where for the most part no special protection is required, it is very widely spread, probably coming next to that of the Fucaceae in the amount of area covered. According to Harvey the plant is washed ashore in quantity durmg autumn; and the hemispherical cushions are collected (under the name of “ figs”) for use as manure (Phye. Brit.). Clare Island Survey—Marine Algae. 15 67 Corallina-Cladostephus Community. This vegetation is frequently very well developed on sandy shores where low rocks are present (¢g., Old Head). It retains a certain amount of sand, but is usually found on raised or sloping rocks where the quantities deposited are small. The community clearly forms a connecting link between the sand- and rock-formations; but as both the species composing it are distinctly saxicolous, and do not form rhizoids, it is regarded as belonging to the latter, and is dealt with on p. 58. The Porphyra umbilicalis Community. Though attached only by a basal disk, and clearly belonging to the rocky- shore formation, P. umbilicalis var. laciniata is frequently found, fringing the sides of half-buried rocks, on sandy beaches. Its growth in this habitat is described on p. 57. Vegetation of Sand-Pools. A very interesting psammophilous flora occurs in pools amongst low rocks on sandy shores. The plants are not small and filamentous, such as Rho- dochorton or Vaucheria (pp. 65 and 64), but large and bushy; hence, though partially embedded in sand, they are invariably fixed to underlying rock. Polyides and Gracilaria may be taken as typical examples; but some six or eight other species occur, all being sand-loving plants, and found as a rule in the sub-littoral region. When, as is often the case, ordinary rock-pools are absent, the constancy of this sand-pool vegetation is very manifest. Algae which thrive under such conditions have, as would be expected, much branched thalli of erect habit. The fronds are more or less terete, and in many cases tough. But other forms occur, especially where the amount of sand is not excessive. Sand-pools are found at Old Head, and between that spot and Leckanvy; also to a certain extent near Mulranny. The typical flora is as follows :— Polyides rotundus, c. Gymnogongrus Griffithsiana, 7. Ahnfeltia plicata, c. Gelidium crinale, r. Gracilaria confervoides, c. Cladostephus spongiosus. Polysiphonia elongata, c. Corallina officinalis. P, atro-rubescens, c. Rhodochorton floridulum. Furcellaria fastigiata. The last three are not characteristic of this vegetation only ; but they are exceedingly common if the sand-layer is not thick, and, as shown above, they form part of the littoral flora of the shore in question. The species of 12 15 68 Proceedings of the Royal Irish Academy. Gymnogongrus and Gelidium were, however, not found outside sand-pools. Where the sides are steep, ordinary saxicolous species naturally occur; but as a rule the sides are shelving and the pools shallow. When fresh water exudes between the rocks, the Rhodophyceae are replaced by Chlorophyceae—i.e., Chactomorpha aerea, Enteromorpha intestinalis, — Cladophora sericea, C. rupestris, and Ulva Lactuca; Gracilaria, however, usually remains, being able to withstand a considerable quantity of fresh water. Where the sand-layer is very thin a number of other species are found, and being characteristic of sandy shores, the most usual amongst them are listed below :— Ulva Lactuca (very thick form). Porphyra umb. laciniata. Enteromorpha Linza. Helminthora divaricata(on Polyides). Monostroma Grevillei (epiphytic). xelidium spp. Seytosiphon lomentarius. Chondrus crispus (broad form). Punctaria latifolia. Gymnogongrus norvegicus (7). Chordaria flagelliformis. Chylocladia kaliformis. Eudesme virescens. Laurencia pinnatifida. Leathesia difformis. L. hybrida. Chorda filum, Polysiphonia nigrescens. Cystoseira granulata. Spermothamnion Turneri (epiphytic). Halidrys siliquosa. Dumontia filiformis. Dictyota dichot. var. intricata. Lithophyllum incrustans. ' (b) Sub-littoral Region. Rhodochorton floridulum Association. As before mentioned, this psammophilous species descends into the sub-littoral region, covering large areas where the bottom is flat and very sheltered, as in the inner part of Clew Bay. The association apparently exists in certain spots as an unmixed growth ; but as arule it is more or less interspersed amongst Lithothamnium or other algae (see p. 65). Zostera Association. The soft bottom of the inner part of Clew Bay is admirably suited to the growth of Zostera, and in the muddy parts especially, extensive beds are found. The characteristic algal epiphytes are also well-developed, being most marked in spring and early summer. Ostenfeld in his report on Zostera (08) points out that, as a general rule, Clare Island Survey— Marine Algae. 15 69 the size of the plant is more dependent on the nature of the bottom than on the depth in which it grows. He shows that the form with short, narrow leaves is found on firm sand, and that the long, broad-leafed form is characteristic of soft mud, rich in food substances. A slight difference between the algal vegetation of the sand and mud Zostera was observed in Clew Bay, but not with sufficient care to warrant comment, except to note that the latter proved the most productive in the way of algae. In spring, masses of Ectocarpi occur, together with innumerable tufts of Cladophora lanosa; whilst im summer and autumn species of Bangia and Chantransia are more abundant. Ascocyclus orbicularis is almost always present, being noted in quantity in spring, summer, and autumn. At the margins of the Zostera-meadows, and in spots where largest ones exist other plants, such as Halidrys, Laminaria saccharina, and Cystoseira granulata, are met with; and the association then tends to approach that which Ostenfeld has termed the “mixed Zostera vegetation ” (l.c., p. 33). Of the larger epiphytes the following are usually frequent in their respective seasons :— Cladophora lanosa. Myriotricha densa. Castagnea Zosterae. Scytosiphon lom. var. zostericola. Punctaria tenuissima. Laurencia obtusa var. crucifera. Desmotrichum undulatum. Polysiphonia violacea. Ectocarpus confervoides. Ceramium strictum. E. siliculosus. Giraudia sphacelarioides, 7. E. sp. Leptonema fasciculatum, *. Litosiphon pusillus. Mesogloia lanosa, 7. Whilst among microscopic species may be mentioned :-— Calothrix confervicola. Melobesia farinosa. Chantransia virgatula. Rivularia atra. C. Thuretii. Isactis plana, r. Ascocyclus orbicularis. Erythrotrichia Bertholdi, 7. Rhodophysema Georgii. The Fruticulose Lithothamnium Association. Probably no group of seaweeds is more generally recognized as forming extensive colonies or associations than the Lithothamnia. Submerged beds, often known as “coral-banks,” occur in the arctic, temperate and tropical regions. In the latter. they are frequently exposed at low water, and play 15 70 Proceedings of the Royal Irish Academy. an important part in the formation of coral reefs. But notwithstanding the many references, both systematic and biological, to this type of vegetation, it is not by any means easy to compare the Lithothamnium flora of one country with that of another. This is owing to the deceptive nature of morphological characters, and to the many erroneous determinations in the past. Foslie has left reliable records for recent years, and the anatomical classification elaborated by Dr. Lemoine promises more success for the future. Between the outer islands of Clew Bay areas of Lithothamniwm caleareum “coral-banks”) are common. They are found on a soft but firm bottom, consisting usually of stones and shells on sand, though occasionally Litho- thamnium may be dredged where there is a thin layer of mud. The usual depth in which the banks occur is 1-5 fathoms, and here and there the plants may be gathered by hand during the lowest tides. The association is com- posed of the following species :— Lithothamnium calcareum, dom. Lithophyllum fasciculatum, Jocally sub-dom. L. incrustans. L. incrustans var, subdichotomum, rare. Lithothamnium norvegicum, rare. In many places Z. calcarewm forms a pure growth extending over a con- siderable area, but in others it is mixed with Z. fascicwlatum, and sometimes all four species occur mixed together. A characteristic growth of epiphytes accompanies the vegetation, of which Polysiphonia subulifera, Pterosiphonia parasitica, Aglaozonia reptans, Spacelaria cirrhosa, Rhodochorton floridulum, and Gelidium sp. are the most constant. Dictyota, Chylocladia kaliformas, and Lauwrencia obtusa are also abundant in summer. With trivial variations this is the type of fruticulose Lithothamnium vegetation found throughout Clew Bay. It presents one remarkable floristic feature, namely, the presence of Z. fasciculatum, a species which is only known from the west of Ireland. Another interesting plant is Z. inerustans, var. subdichotomum, an alga which was formerly regarded as L. dentatum, but shown by Dr. Lemoine in the present report to be a detached and rolling form of the common species, Z, inerustans. Other Lithothamnium banks in Ireland and Great Britain.—It was of interest to find that the Clew Bay association agreed exactly in its composition with that which had been described for Roundstone. The latter district is the original station for Z. fasciculatwm ; and Foslie, when working out the European Lithothamnia, paid a special visit to the locality and Clare Island Survey—Marine Algae. 15 71 published an account of his trip (99). He found Z. fasciculatum still plentiful, as it was also in September, 1911. The dominance of ZL. calearewm in both Roundstone and Clew Bay is normal for the British Isles. The remarkable “coral beaches” at Mannin Bay and Ballyconneely Bay (Co. Galway) are due to the presence of extensive Lithothamnium associations in the neighbourhood. When driving from Clifden to Roundstone in September, 1911, the former spot was examined; but time did not permit of exploration by dredging. It was evident, however, that the material washed up is Z. caleareum, and that the beach, a fairly wide one, is composed almost entirely of broken particles of this species. The photographs on Plate IV, kindly taken by Dr. G. H. Pethybridge, show two views of the beaches in Mannin Bay. A few low rocks occur in part of the bay as shown in the lower illustration, but elsewhere there are stretches of pure “coral sand.” The greater part of the sand is composed of small particles (see Plate V, fig. 1); but at high-water line and in certain other places coarser pieces and knolls accumulate, with a certain quantity of broken shells. Dr. Pethybridge informs me that at low-water mark the particles are exceedingly fine and resemble ordinary sand ; but it is probable that they have the same origin. he Lithothamnium fronds are completely bleached, and the beach is thus dazzlingly white in the sun. ‘Though the coral sand is carted for use as manure, fresh supplies are constantly washed up, so that it is evident that the bank outside is extensive. A similar beach is said to exist in Bantry Bay.! In the British Isles LZ. calearewm is certainly the dominant fruticulose species, beds being known from several localities (Studland and Falmouth in the south of England, and Bute in Scotland), and under certain conditions it is gvashed ashore in quantities. A fine bed in 14 fathoms was once dredged by me off Fowey (Cornwall). Another species, Z. racemus, appears to exist as an association at Falmouth, since it is washed ashore in plenty; but this is the only station for the plant in England, and records of Lithothamnium banks in England or France formed by species other than ZL. calcarewm require verification, Comparison with other countries.—L. caleareum is also frequent in France. It is abundant in Normandy, and in Brittany as far south as Croisic. Lemoine (11) states that it is known from Denmark, Portugal, Naples, Morocco, and Algeria. In all these localities it probably occurs in associations similar to 1 Several other remarkable beaches occur in the west of Ireland. A shore consisting exclusively of broken shells is found at Knockboy (Co. Galway), which, like the coral-sand beaches, is exploited for manure (Johnson and Hensman, ’99); whilst Dog’s Bay, near Roundstone, is famous for its beautiful sands, composed almost entirely (up to 90 per cent.) of Foraminifera, 15 72 Proceedings of the Royal Irish Academy. these found in the British Isles. At Morlaix and Concarneau there is “coral sand” or “Maerl,” and, as in Ireland, this is used for manure. Lemoine (710) has also published a detailed account of the biology of the plant. She notes that at Concarneau the bottom of the bay is practically covered with L.. caleareum, and fragments of living fronds are continually cast up by the — waves. Several “coral beaches” occur on the islands of the Glenan Archipelago, but there, as at Mannin Bay, the fragments washed ashore are mostly dead. In the Arctic seas, Kjellmann writes of large areas, in 10 to 20 fathoms, covered with Lithothamnia, of which LZ. glaciale and L. soriferwm are abundant. At Mussel Bay in Spitzbergen, Z. glaciale is said to cover the bottom to an extent of 4 to 5 square miles; whilst off the north of Norway and on the coast of Iceland, Z. Ungeri is abundant. In the former country banks of L. norvegicum also occur, The association is represented in Greenland by L. Ungeri. In the south of Norway the floristic features are more in agree- ment with those of Britain, and Z. calcarewm becomes prominent. An association composed largely of this species has been described by Gran for Kristianiafiord (793). Reference must be made (though it is outside our usual circle of com- parison) to Madame Weber van Bosse’s account of the Lithothamnia of the “Siboga” Expedition (04). A very luxuriant vegetation exists in the shallows and reefs of the Malay Archipelago, and several communities on different types of ground are referred to. A description is given of a locality where the Lithothamnium knolls are rolled backwards and forwards between the islands by the tidal current, whilst in other parts of the same channel they are heaped up in banks. Two photographs are reproduced of an extensive bank of LZ. erubescens var. Haingsisiana, which Madame Weber describes as being “covered as far as the eye can reach by the pretty, beautifully pink-coloured knolls, which are heaped so close together, that while walking one crushes them continually, making a peculiar noise as of broken china” (/.c., p. 5). Geological Importance—In localities where masses of Lithothamnium accumulate on the sea-bottom, these algae play an important part in rock- formation, and masses of limestone containing well-preserved Lithothamnia occur in various geological strata. Structureless limestone also may be derived from the same source, for Walther, working at Naples, showed that the structure of recent Lithothamnium nodules is gradually obliterated by the action of percolating water. The literature on the subject is scattered; but Seward’s paper (94) gives a general summary with a full bibliography ; and a brief but up-to-date account has just been published by Lemoine (ale) Clare Islund Survey—Marine Algae. 15 73 The Pebble-attached Association of Qumet Bays. This association represents a definite type of sub-littoral vegetation, which is found on a soft bottom strewn with stones and pebbles. A very large number of species go to form it, and they are for the most part characterized by finely divided bushy fronds, though heavier and less bushy plants are found on the larger stones and on the occasional pieces of rock. In some localities almost every stone down to the smallest pebble bears a tuft of one or more algae. The latter consist of members of the Green, Brown, and Red groups; and, though they vary considerably in different localities, the mixture of colour is a marked feature and suggests comparison with Kjellmann’s “ Bunte Formation.” But both the constituents of that association, and the depth at which it oceurs, are different from those here described. The present vegeta- tion corresponds rather to Borgesen’s Stictyosiphon association and to Boye’s Dictyosiphon-Spermatochnus-Corallina formation, though Dictyosiphon is not always present. The species are annual or short-lived, which in part accounts for the varied aspect presented during the season. The association extends from just above low-water mark down into 3 or 4 fathoms. Its shallower regions may be studied in detail by wading at spring tides, when the bushy fronds of Polysiphonia, Cladophora, Spermatochnus, Dictyosiphon, ete., may be examined im situ. Dredging in deeper water shows a very similar type of growth, though some species disappear and others become more abundant. The colour of the red species improves with increased depth. The association, which reaches its maximum development in May, is well seen in many of the inner parts of Clew Bay, and has been studied specially on the west side of Annagh Island, and, in a slightly modified form, in Bellacragher Bay. In brackish localities an increase of Chlorophyceae is apparent. «\s it oceurs in Clew Bay the principal species are as follows :— (2) In spring — Enteromorpha clathrata. Lomentaria clavellosa. Cladophora Rudolphiana. Nitophyllum punctatum, Striaria attenuata. Polysiphonia urceolata. Stictyosiphon subarticulatus. P. elongata. Dictyotosiphon foeniculacea. P. nigrescens. Ectocarpus siliculosus. P. atro-rubescens. Pylaiella littoralis. Ceramium rubrum. Cystoclonium purpurascens. Griffithsia corallina. Delesseria hypoglossum. R.I.A. PROC., VOL. XXXL, K 15 15 74 Proceedings of the Royal Irish Academy. (6) In summer Striaria, Dictyosiphun, Delesseria, and Griffithsia largely disappear, but the other species remain; and in addition we find :— Cladophora rectangularis. Polysiphonia subulifera. fo) Spermatochnus paradoxus. P. violacea. Castagnea virescens (shallow P. fibrillosa. water). Bronegniartella byssoides. fo) Dictyota dichotoma. Ceramium spp. As autumn advances one species after another becomes detached and disappears, though Dictyosiphon, Ectocarpus, and Polysiphonia nigrescens remain. 7 NW Sea | 1) o in ~ r tA PROC.R.TACAD. Von. XXXI. PART. 15; Pi AT) 1X. LTS J lo ? pl a= r wee TT} © 4 LE \) i ea West, Newman lith Del. A. D.C. SUBVEY,— COTTON: MARINE ALGAE. tilothamnion lucifugum, sp. nor. ARE ISLAND j C XG PART 15, PLATE Solaigea = c=) Uae Qo on Goo Oa ue WOGBt * West,Newman lith. 1-3. Galothrix endophytica sp. nor. 4-9. Ascocyclus Saccharinae, sp. nor. N. laceratum var.uncinatum Grer. i. J, Ag. 10. Nitophyllum uncinatum , LAND SURVEY.- COTTON: MARINE ALGAE. Qa 2) CLARE Is PROC. R.I.ACAD. Vou. XXXI. et ea Ou aS A a PAOD rial iC a Del. A.D.C. PLATE XI, 5 PART 1 NXXI. LAGAD., VoL. R. Proc, Saddle Head ga Has Dooe 8 MILES. . SCALE. Map of the Clare Island District. MARINE ALGAE. CLARE IsLAND SURVEY.—COTTON : SMITHSONIAN INSTITUTION LIBRARIES (im =r ae ee Te ~ ROYAL IRISH ACADEMY CLARE ISLAND SURVEY SCHEME OF PUBLICATION — ay [The parts already published are marked with an asterisk. The whole willform : ** * (© 00 —3 G> cn Wy CO BD * 10 * 41 3 3k tn % 14 3 15 % 16 % 17 % 18 *% 19 #20 2 # 22 #28 % 24 25 % 26 27 28 % 29 30 31 Vol. XXXI of the Proceedings of the Academy. ] Grnerat Description, anp Narrative. BR. Ll. Praeger. ane History anp Arcuaronocy. T.J. Westropp. 78 pp. 10 plates. 4s. Prace Names. John MacNeill. T. J. Westropp. Gartio Puant anp Antuan Nawms. N. Colgan. 380 pp. 6d. Acricutture. Prof. James Wilson. 46 pp. Is. Mersorotocy. W. J. Lyons. Guotoey. J. R. Kilroe. TT. 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Hewmerera, J, N. Halbert. 10 pp. Oxrnorters, Prof. G, H. Carpenter. 4 pp: ; [For continuation see page 4. | 6a, i: om 2B rok: ed H. Carpenter. Hs Ananerpa. 'D, R, Pack Beresford. 8 pp Puaraneipa. D. RB. Pack Beresford. 2 PP. Anoriscowa. James Murray. 16 pp., 8 plates) a Psevposcorerones. H. Wallis Kew. 2pp. § %& 391 Acarmuna: i—Hypracarmwa. J.N, Halbert. 39ii Orneg Acsnmipa. J.N. Halbert. — % 40 Descarops. G.P.Farran. Spp. Nell ai % 441 Sonrzoropa anp Cumacua. 'W. M. Tattersall. 10 pp. ' 2 “Ampmrropa. W. M. Tattersall. * 43 Isopopa—Marmn. W. M. Tattersall. 6 pp. 1 Ga. %* 4% Tsoropa—Trrnesrran. N. H. Foster. ‘4 pp.’ 4§ Ewromosrraca—Maning. G, P. Farran. % 46 Enromosraaca—Fresuwares. D. J. Scourfield, 18. pp. 1 plate Potyowanrs anp Ancurannenipa. R. Southern. Oxicoonsrrs. BR. Southern. -Guenvrea, RB. Southern. Himvowea, BR. Southern. ‘Rommera (excluding Bozxroma). OC. F ‘Ronaselot.. 10 a) Rorirera Borinoma, James Murray. 20 pp. 1 igs Ponyzos, A. R. 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