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