Littoral ecology of Marion and Prince Edward Islands (Southern Ocean)

Anthonie F. de Villiers

Zoology Department University of Port Elizabeth P.O. Box 1600, Port Elizabeth 6000

The shores of the volcanic islands Marion and Prince Edward Die kuslyne van die twee vulkaniese eilande Marion en are rocky and consist mainly of steep cliffs and beaches Prince £dward is rotsagtig en bestaan hoofsaaklik uit kranse composed of boulders the sizes of which vary from a few en strande gevorm deur rotse wat kan wissel in deursnee van centimetres to several metres in diameter. All the shores are 'n paar sentimeter tot etlike meter. Hoewe/ a/ die strande exposed, those with a westerly aspect being severely so. blootgestel is, geld dit vera/ vir die met 'n westelike aansig. Tidal range is 71 cm at springs, 21 cm at neaps. Swell Getyhoogtes wissel van 71 cm met spring, tot 21 cm met heights on the east coast vary between 0 and 4 m. Salinity dooiegety. Die hoogte van deinings aan die ooskus wissel and temperature are typical of the area. Gales, high humidity, tussen 0 en 4 m. Soutgehalte en temperatuur is tiperend van abundant precipitation and relatively little fluctuation of die gebied. Die klimaat van Marion en Prince Edward word temperature characterize the climate. Basic zonation on gekenmerk deur stormsterktewinde, hoe humiditeit, oorma­ plane sloping rocky faces consists of a black lichen tige presipitasie en relatief min wisseling in temperatuur. (Verrucaria) zone in the supra-littoral, a Porphyra zone in the Basiese sonasie op gelykver/opende rotsaansigte bestaan uit mid and upper littoral, a kelp zone of Durvillea antarctica in 'n swart ligeen Verrucaria sone in diesupra-littorale gebied, the lower littoral, and a lower red zone of delicate 'n Porphyra-sone in die middel en hoer littorale gebied, 'n rhodophytes in the upper sub-littoral. kelpsone bestaande uit Durvillea antarctica in die laer Five transacts on different types of shore showed up littorale gebied en 'n /aer rooisone van delikate Rhodophyta departures from the basic pattern as a result of increased in die boonste sub-littora/e gebied. shelter, increased shade and decreased stability of substrata. Vyf transekte Op verskillende tipes van kus/yn het The shallow sub-littoral is characterized by abundant afwykings van d;e basiese patroon getoon a.g.v. 'n toename corallines and other rhodophytes as well as phaeophytes of in beskutting en .skaduwee en 'n atname in die stabiliteit van the genera Desmarestia and Durvillea. In parts large die substraat. Die v/ak sub-littorale gebied word gekenmerk aggregations of the patellid Nace//a delesserti were found. deur welige Corallina en ander Rhodophyta asook Phaeo­ Pools are infrequent and biota in them vary with salinity. A phyta van die genera Desmarestia en Durvillea. Plek-plek is wide variety of invertebrates are associated with the kelps groot getalle van die patella Nacella delesserti aangetref. Macrocystis and Durvi/lea. Trophic levels consist of primary Poele kom se/de voor en die biota daarin wissel volgens producers and primary, secondary and tertiary consumers. soutgehalte. 'n Groot verskeidenheid van Invertebrata kom Zonation conforms with the pattern noted on other sub­ op die kelpe Macrocystis en Durvillea voor. Die trofiese Antarctic islands except Tristan da Cunha. lt is not possible vlakke bestaan uit primere produsente en prim§re, sekondere to adapt the universal scheme of zonation of Stephenson & en tersi§re verbruikers. Stephenson (1949) effectively to the pattern on Marion and Sonasie j(om ooreen met die patroon wat op ander sub­ Prince Edward. The biogeographical affinities lie within the Antarktiese eilande waargeneem is, met die uitsondering van sub-Antarctic and to a lesser degree within the Antarctic. The Tristan da Cunha. Dit is nie moontlik om die universe/e young age of the islands and relatively recent glacial ana skema van Stephenson & Stephenson (1949) effektief aan te volcanic activity may account for paucity of species and low pas by die patroon wat op Marion en Prince Edward voorkom endemicity. nie. Die biogeografiese ooreenkomste is tipies sub­ Antarkties en tot 'n mindere mate Antarkties. Die beperkte diversiteit van spesies en die gebrek aan endemiese spesies kan moontlik togeskryf word aan die betreklike jong ouderdom van die eilande en relatief resente aktiwiteit van gletsers en vulkane.

INTRODUCTION Marion Island (46°54'S, 37°45'E) and her smallet Marion reaches an altitude of I 230 metres and is neighbour Prince Edward (46° 38'$, 37° 57' E) are 290 square kilometres in extent. The smaller Prince situated in the southern Indian Ocean 2300 Edward, lying 22 kilometres north-north-east of kilometres south-east of Cape Town. Arising about Marion, has an area of 44 square kilometres and 370 kilometres south-east of the crest of the Mid­ reaches an altitude of 672 metres. Indian Ocean Ridge, they are considered to represent Annexed by South Africa in 194 7/ 48, the islands the summits of coalescing shield volcanoes ( Verwoerd, J first became the site of extensive scientific research 1971). The oldest lavas, recognisable in the field by when Professor E.M. van Zinderen Bakker of the their grey and generally smooth appearance have a University of the Orange Free State led the first dated age of about 276 000 to I 00 000 years, while the South African Biological/ Geological Expedition younger lavas which are black and rough were there during the summer of 1965/ 66. The expedition extruded about 15 000 years ago. (McDouga/1,2 enjoyed the support of the South African Scientific 1971 ). Committee for Antarctic Research (SASCAR) and S. Afr. J. Antarct. Res., 1976, Suppl. 1 2 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 3

0 was sponsored by the Department of Transport. The 60 300 0 not been studied and appear to have been lost. I . o six-man team investigated the avian ecology, botany, Identifications and ecological notes on the following ,..,.;..,.:,SOUTH. AMERICA .,1 \ .. fTr;chn n .. runh" t . 1300 palynology, marine biology, geology and topogra­ groups are recorded in the 1971 monograph: Pisces, '•:.,.._ ...... , phy. Many organisms were collected and distributed Echinodermata, Holothuroidea, Mollusca, Oribatei, ...... ' AFf:liCA to specialists for identification. Entomostraca, lsopoda, Polychaeta, Hirudinea, ..J The results of the expedition were published in Hydrozoa and Actiniaria. A paper by Mitchell­ \ 1971 in an extensive monograph to which some 40 Innes14 (1967) entitled "Primary production studies in ' authors contributed articles (Van Zinderen Bakker et '·-, the South-West Indian Ocean 1961-63" dealt with the \ \ a/.,3 1971). This major work laid the foundation for Africana 11 autumn and winter cruises to Marion and '), ?-. the more intensive investigations during the summer the Crozet islands during the International Indian ., , \ expedition of 1971 / 1972 which was geared towards Ocean Expedition. 900 ',,...... _ ,. Pr1~ce Edward I. obtaining insight into the bio-energetics of the Nef1S ( 1968) worked on the microplankton \Ma rio~ l· island's ecosystems. The limnology of streams, lakes gathered during this cruise, including diatom \ ' \ \ \ and tarns, the role of nitrogen and the cycling of distribution in the sea around Marion and Prince \ :· Is .• Crozet Edward. Sawyer16 ( 1972) recorded a new leech I • minerals was studied. In spite of these advances a 1 I 60° \ : serious gap in knowledge of the island's ecology still (Annelida : Hirudinea) parasitic on a nototheniid fish \ existed in 1972, since no full account of the ecology of from Marion and the Crozet islands, and Sawyer and de Villiers (in press) have presented further I \ rocky shore organisms had been published. The u., jiS. Kerguelen author was appointed as marine biologist to the third information on littoral leeches. Recent publications (J. I I ~· I ; I expedition (during the summer of 1972/73) to stemming from the 1972/73 collections include ~ / I 0 ~~ I • investigate this aspect. information on oribatid mites (Acari) (Engelbrecht, 17 40 - ) V I ' I "" I I I 1~ I I 1974) , echinoderms ( Rowe & Clarke, t8 1975) and 120° I \ ~/ isopods (Kensley, 19 1975). ·,_ \ ',,, ~ / I Previous marine biological investigation at Marion ·,,, vv and Prince Edward islands Literature on sub-Antarctic littoral ecology ·., "'\., ~~ ' v" / 90° The island was discovered in 1772 . by the Fr·ench In this paper the author has followed the sub­ ·., .. ~ / seaman Marion Dufresne, but the first opportunity Antarctic region described by Deff20 (1972). It ~~~ / ...... f>. /'/ for scientific study arose a century later when, on 26 includes southern South America from Tierra del ' ...... ___ ...\ _____ ....---- Fuego to about latitude 47°S on the west and east ·-...... December 1873, a small party was able to go as.hore ·.. ,, coasts, as well as the following islands: Falklands, on Marion's north-east coast during the Challenger's ' investigation of the sub-Antarctic. Among those Tristan da Cunha group, Marion and Prince Edward, ··,··-;.., present was the naturalist H.N. Moseley who made Crozet group, Kerguelen, Macquarie, Auckland the first littoral observation on Marion. He drew islands and Campbell Island (Fig. 1). attention to the large brown alga Durvilleaand noted Publications relating specifically to littoral ecology that it played a marked role in breaking the surf in the Sub-antarctic are few. Special mention, 150° (Moseley, 4 1879). With only a few hours at his however, must be made of Knox who has produced W 180° E 120° disposal, Moseley concentrated on the terrestrial some most useful summaries of zonation patterns on environment and did not collect organisms from the the shores of sub-Antarctic islands (Knox,2t 22 23 24 Fig. I. The positions of the sub-Antarctic islands (after Defl. 1972). shore. H e did however, collect several algae from 40 1960; 1963; 1968; 1970). fathoms while aboard the Challenger (Dickie,s 1877). Valuable information on the distribution and species there. Isobel Bennett'sJ6 "Shores of Macqua­ habitats were then selected for intensive study. These Collections of invertebrates and fishes were also ecology of algae on the shores and in the sub-littoral rie Island" ( 1971) provides a popular account of the areas were of necessity all on the relatively sheltered made while the Challenger was in the vicinity of the of sub-Antarctic South America and the Falkland littoral zone with many useful photographs. Littoral east coast, because of the inaccessibility of the west islands. Further collections were made by dredging islands may be obtained from Skottsberg2s (1941). zonation on New Zealand's sub-Antarctic islands has coast and the difficulty of working there. The areas from Discovery 11 in 1935 (Antarctic Pilot,6 1961}. In The zonation of algae and invertebrates on the shores been briefly described by Knox31 (1963) and M orton included Ship's Cove, Duikers P oint, Transvaal Cove 1939 a party from the Bougainville landed briefly on of southern Chile as far as latitude 54° S have been & Miller38 (1968). and Macaroni Bay (Fig. 2). Marion's north coast. Rene J eannel, an entomologist documented by Straehf26 ( 1964), and there are brief from the Paris Museum and the only naturalist notes in Stephenson & Stephenson21 ( 1972). MATERIALS AND METHODS aboard, collected marine organisms but these have Baardseth28 (1941) has given an account of the Surveying the selected areas never been studied and are presumed lost (Jeanne/, 7 algae of Tristan da Cunha, while the algae of Gough Physical data The slopes of four shores (Ship's Cove, Duikers 29 Point, Transvaal Cove, Macaroni Bay transect 2) 1941 ). have been studied by Chamberlain ( 1965). Both of Physical parameters such as climate, salinity, currents from the upper supra-littoral to the upper sub-littoral Soon after the establishment of a weather station these workers include important ecological observa­ and sea temperatures have been well documented in (in the sense of 42 1960) were measured along on Marion in 1948, biological observations by South tions in their otherwise primarily taxonomic papers. the past and were given limited attention during the Knox, 40 transect lines running normal to the shore with Africans began to appear (e.g. Bennets,s 1948; The littoral ecology of Gough Island was studied present survey (see Schultze, 39 1971; Deacon, 1937; Fuller,4 1 1967). Tidal data were however very limited surveyors' ranging rods according to the method of Crawford, 9 1952; Rand, IO I 954; A Is ton and Schelpe, 11 during the 1955/ 56 Gough Island Scientific Survey 1957; La Grange,l2 1962). These papers were confined and written up as a traditional rocky shore intertidal and several attempts were made to establish the tidal Dayn (1959). The slope of a very steep shore to terrestrial life. investigation (Chamberlain, Ho/dgate & Wace30, in regime. These included the construction and installa­ (Macaroni Bay transect I) was measured using a tion of an automatic continuous tide recorder, and spirit level. In 1967 N.R. Fuller,13 marine biologist to the first preparation). Zonation of algae on the French islands South African Biologjcal and Geological Expedition Crozet and Kerguelen has been documented by the taking of readings from a graduated rod. Algae and invertebrates were collected one metre 32 to Marion and Prince Edward islands (during Delepine & Hureau31 1968) and Delepine ( 1966). on either side of the transect lines and careful notes 1965166) published a preliminary account of the The ecology of littoral invertebrates on the shores of Selecting representative shores on their position and relative abundance were shores (since many of the organisms he had collected Kerguelen Island has been described by ArnaudJ3 Field trips were made to various points on the island recorded. The organisms were preserved in alcohol or awaited identification). The limited information (1974). in order to gain a broad impression of conditions and formalin for indentification. the way in which organisms varied with them. Such available led to misinterpretation of the zonation Kenny & Haysom34 (1962) described the general The transects were carried out over a period of 3 pattern on Marion's shores. While many of Fuller's ecology of rocky shore organisms at Macquarie while trips, often of several days duration, were made on months (January to March) and it is possible that foot, so that scientific equipment had to be kept to a comprehensive collections were lat~r identified and Simpson~s ( 1976) has presente.d detail~d in_formation seasonality of certain algae may have biased the results published, some important groups have on physical and biotic factors mfluencmg SlX mollusc minimum. Areas representative of the major littoral comparisons. This fact should receive attention in the 4 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 5

(b) Sloping unbroken rock: This is an uncommon Slope and localized topography type of shore on Marion but is easy to survey and Marion Island's unbroken shores are primarily made collect on. The basic pattern of sub-Antarctic up of vertical or very steep cliffs and it is rare to find shores of considerable extent with an even slope zonation described by Knox45 ( 1968) is well illustrated on this type of shore at Macaroni Bay. inclined at less than 50° to the horizontal. A notable t exception occurs at the northern extremity of TRUE NORTH (c) Low reefs: These extend seaward as a series of low (often flat) rocky shelves and are frequently awash. Macaroni Bay. They often occur at the base of cliffs. Examples may Broken shores, on the other hand, may slope gently be found at Mixed Pickle Cove, 2 km west of Cape to the sea as boulder beaches (Transvaal Cove) or Triegaardt Bay Hooker, and at Transvaal Cove. steeply as chaotically piled blocks of lava (eastern (d) Broken shores: Marion's sea cliffs are frequently extremity of Crawford Bay). Intermediate conditions interrupted by boulder-lined shores. The boulders between these extremes occur at various points. may occur as a narrow strip along the base of the Where sea cliffs have collapsed (immediately cliffs, (south of Kildalkey), as boulder beaches, south-west of Kildalkey Bay) a new shore of lesser Cape (Transvaal Cove) or as chaotically piled steeply slope is created, resulting in increased available sloping shores (eastern extremity of Crawford Bay to surface area for colonization. In addition the Cape Hooker). fragmentation of what was formerly a two­ A gradation in shape and texture of the boulders dimensional face results in a multi-dimensional shore occurs with size and degree of exposure. ·where and an increase in habitat diversity. Apart from the smallest (less than IOcm diameter) they are rounded overall slope of the newly created shore, there are the Soft Plume River and polished and form stony beaches such as at individual slopes of the resulting boulders, a decrease Trypot Beach and Kildalkey Bay. Boulders of about in light penetration, exposure and particle size, and 30 - 100 cm diameter as found south of Soft Plume an accumulation of detritus beneath the boulders. River are also round, but as a result of increased Topography at any one point influences the stability are covered by lichens and algae. Very large effective tidal range by either damping the swell or in boulders several metres in diameter are frequently some cases augmenting it. angular and of rough texture and bear rich growths of intertidal organisms. Such shores may be found at Aspect and exposure Ship's Cove and Cape Hooker. Oceanic islands afford excellent opportunity for the study of the effects of aspect since shores facing every KILOMETRES A comparison of boulders of similar size on the west and east coasts illustrates the consistently severe point of the compass occur yet are all situated in the Fig. 2. Outline of Marion Island showing localities mentioned in the text. Solid circles indicate transect sites. wave action on the west coast, since the boulders same water mass and climate. there are generally less angular and have a sparser Lewis41 ( 1972) has pointed out that "extreme covering of lichens and algae. exposure" quoted by one worker may be only future. Colour photographs of the selected shores RESULTS (e) Sand or shingle beaches: these are rare and are "moderate exposure" to another with wider experience. were taken and later used to check the levels reached Standardization of terms on a world-wide basis has not by characteristic organisms. periodically eroded away by storms to expose underlying boulder shores. A sandy beach at Ship's been possible primarily for two reasons. Firstly, no apparatus has yet been devised which can be used to Study of shores distant from base camp Cove is composed of fine black volcanic particles and The littoral environment of Marion measure all the physical factors which together Several trips to the very exposed west, south and is the only one of its kind on Marion. South-west of constitute "exposure" in its broadest sense. Secondly, south-east coasts were made. Severe wave actio n in Island (physical aspects) Kildalkey Bay a very small cove floored by fine The nature of the coast shingles was encountered while on the eastern shores biological indicator species are geographically re­ these regions precluded detailed sampling and 4 surveying. Levels reached by zone-forming organisms Marion Island's 72-km coastline is roughly oval in of Goodhope Bay a shingle beach may be found. stricted to limited areas (Ballantine. 8 1961 ). were estimated from adjacent heights using binocu­ outline (Fig. 2) and owes its sculpturing to the Organisms living on the west coast of Marion lars. Colour photographs of these shores served as a competi tive processes of volcanic extrusion and Substrata Island must face exposure equivalent at least to the useful check. Organisms were collected wherever marine erosion ( Verwoerd, 44 1971 ). Conclusive evidence of glaciation on Marion Island is most severe in the world. The wind, which usually has possible but this often involved brief dashes into the In the case of the lavas of the older grey succession provided by the occurrence of striated platforms on a westerly component, accelerates to gale force on littoral between successive swells. Localities investi­ the original shores have receded to such an extent the older grey lava as low as 140 metres above sea more than 100 days a year and may blow over a fetch gated included Cape Davis, Mixed Pickle Cove, that 30-m high cliffs plunging directly into the sea level ( Verwoerd, 40 1971 ). It is likely that the of 8 000 km. There are virtually no offshore rocks or Goodhope Bay, Rook's Bay and the Cape Hooker (King Bird Head) have been formed. Elsewhere smoothing action of the ice occurred at sea level also. reefs to modify the strength of the swells and for the area. marine erosion has resulted in the smoothing out of The younger black lavas which were extruded after most part they travel unimpeded in deep water to initial irregularities and many shallow, gently curved the glaciers receded are generally of a far rougher expend their energy against the sea cliffs. Investigation of the shallow sub-littoral bays now occur (Macaroni Bay). In contrast the texture than the old grey lavas. Both types of lava The land mass of Marion Island refracts the Since many organisms occurring in the littoral are younger black lavas terminate in the sea as a series of have been subjected to normal weathering by the prevailing swell such that the north and south coasts basically outliers of sub-littoral communities, it was irregular inlets, capes, off-shore rocks, stacks, arches force of waves, abrasion by sand and grit and by are also battered by heavy seas though not as decided to extend the shore survey at Transvaal Cove and caves. rotation of boulders against each other. The action of severely as the west coast. By the time the east coast 3,5 m into the sub-littoral by snorkel-diving. The Nowhere on Marion does a headland or bay attain foot, flipper and body of the great number of animals has been reached most of the swell's energy has been slope of the bottom was measured at intervals along a such proportion that a sheltered shore is created, and that come ashore at favourable points has also dissipated and the calmest conditions generally occur weighted line by two divers. Organisms were collected the entire littoral environment may be considered contributed markedly to the wearing down of the between Duiker's Point and Macaroni Bay. along the transect, but it was not possible to do this in exposed. substrate. the same detail as in the littoral. Several types of rocky shore can be recognized: Grooves, clefts, depressions and ridges occur on Tides General collections of sub-littoral organisms were both types of lava and play an important role in the Several attempts have been made in the past to made in the vicinity of Transvaal Cove. (a) Vertical cliffs: These are from a few metres to 30 m distribution of certain organisms. Bones, feathers, establish the tidal regime on Marion Island (Fuller, 49 in height and dominate much of Marion's coastline. crushed shells, seaward fragments and grit may 1967; Langenegger & Verwoerd,SO 1971; Crawford Prince Edward Island Good examples occur at King Bird Head, Cape accumulate under boulders as mixed detritus. This pers. comm.) Fuller, Crawford and the author all Data were gathered during a one-week visit to Prince Davis, and the Triegaard Bay and Goodhope Bay type of substrate harbours many forms of invertebrate took readings from a graduated rod and found values Edward Island in April 1973. areas. life and is the richest in species. of 76-91, 66 and 70 cm respectively for normal spring 6 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 7

range. An automatic tide recorder used during the indicators may be seen in the transect profiles and are temperatures. While the actual heights obtained may Table 2 1972{73 expedition was destroyed by a storm shortly be inaccurate, the consistency of their observations also discussed on p. 26. Terminology used is that of after installation. enabled the author to check levels reached by high Salinity and temperature in three pools at Transvaal Cove Knox72 73 (1960, 1968). In November 1974 the South African Navy and low tides on days of average swell conditions. In installed a pressure-sensitive tide gauge near the this way the effective tidal heights at Transvaal Cove 0 Temperature (0 C) Salinity / 00 MACARONI BAY TRANSECT I could be assessed. It is stressed that this data is meteorological station. The pressure-sensitive element Max M in Mean Max Min Mean This is one of two transects surveyed at the northern relevant to Transvaal Cove only. was situated 219 cm below mean sea level and gave extremity of Macaroni Bay some 2 km south-east of consecutive readings from November to Analysis of one year's data (March 1972 to 36,0 30,0 33,1 !U! 7.8 17 25 Pool I 7.0 Transvaal Cove. The smooth lava of the older grey February 1973) of swell hei$!:hts has shown that swells Pool 2 37,0 29,3 33, 1 17.0 7.0 9,8 November before being shifted by very heavy swell. At succession slopes steeply down to the sea. Four zones the time of the moon's first quarter the minimum of 0 or 0,5 m occur 67% of the time; I or I ,5 m 24% of Pool 3 38,0 6,0 31,1 16.5 7.0 9,9 ranging from the lower supra-littoral to the upper the time; 2 or 2,5 m 6% of the time; 3 m or greater 3% range was 21 cm and halfway between observed neap sub-littoral can be distinguished on this shore (Fig. 3; tides and presumed spring tides a range of 46 cm was of the time. complex of rocky outcrops known as Paddy Rocks Plates I and 2) obtained. Assuming a roughly sinusoidal tidal range An important factor is the periodicity of extreme extend seaward irregularly and a few pools occur curve, the spring range on 14 November would have conditions such as violent storms or prolonged among them. One is connected to the sea by a lava l. Lichen zone: This zone is predominantly dark been 46 + (46 - 21) = 71 cm. periods of exceptional calm. Ii was found that swells tunnel (Pool I), one receives splash with every tidal grey to black in appearance as a result of the The tidal readings obtained by the South African attained 2 m or higher at least once a month cycle (Pool 2) and one is occasionally reached by continuous encrustation of lichens of the genus Navy are the most reliable available and the data throughout the year except in November 1972 and splash (Pool 3). Verrucaria. It extends from the supra-littoral to obtained by Fuller5 1, Crawford and the author lend February 1973. Swell heights of 4 m or greater Salinity and temperature were recorded in the three HWS where it becomes obscured by the general support to the values obtained. For the occurred once in April, once in September and three pools on 24 occasions between 22 February and 26 rhodophyte Porphyra. purpose of this report the naval prediction of 71 cm times in June. Conversely the longest period during March. Table 2 gives the most relevant findings. 2. P orphyra zone:Porphyra grows luxuriantly from range for springs and 21 cm for neaps will be used. It which the swell height was less than 0,5 m occurred in HWS in the upper littoral to near L WN in the is interesting to compare the spring range on Marion February and lasted for 8 days. In October and Climate mid-littoral where it thins and shares the rock face with the spring ranges on other islands of comparable November two periods of 6 days occurred when the Schultze66 (1971) has presented comprehensive data with light growths of Enreromorpha bulbosa. latitude. swell was less than 0,5 m. taken from more than a decade's records. The climate 3. Kelp zone: The upper limit of the holdfasts of the is characterized by an almost continuous succession fucoid Durv illea antarctica is clearly defined and of extra-tropical cyclones causing a high degree of forms a natural barrier at approximately LWN. Range at H ydrology of coastal waters The shores of Marion and Prince Edward Islands are cloudiness, abundant precipitation and frequent Durvi//ea dominates the shore almost to L WS in U>cality Spring Tide Reference winds which reach gale force on average on 107 days (cm) bathed by sub-Antarctic surface water which flows the lower littoral slightly north of east as the great West Wind Drift per year (10 years' records). Winds with an easterly 4. Lower red zone: Delicate rhodophytes of the Macquarie 137 Kenny and Haysom,sz between the Subtropical Convergence and the component are rare. Thus the north-west, west and genera Schizoseris and Po/ysiphonia dominate 1962 Antarctic Convergence. However, shallow ridges south-west coasts of Marion are lee shores and the upper sub-littoral from LWS downwards. Port Christmas, Kergue- 110 Delepine and Hureau,sJ between Marion and Crozet cause a northward experience very severe wave action throughout most They partly conceal a continuous cover of Ien 1966 movement of water in the vicinity of these islands of the year. While the size of the waves raises effective lithothamnia. Port-aux-Francais, Ker- 160 Delepine and Hureau,s• (Deacon,flO 1937). tidal heights by several metres, the wind itself may guelen 1966 Marion and Prince Edward lie about km north bodily lift the crests off breakers and bear them Details of the zones Gough 120 Chamberlain Holgate & 160 of the mean position of the Antarctic Convergence, inland in the form of coarse splash or spray. Lichen zone: Scattered growths of Prasio/a crispa Wace,ss (In prep.) Huntley61 (1967) has drawn attention to the mark the upper limit of this transect. The generally East Falkland 210 Skottsberg,s6 1941 but sea surface temperatures of less than 3° C are not exceptionally high index of oceanicity enjoyed by smooth lichen-covered rock face is exposed to the West Falkland 425 Skottsberg,ll 1941 uncommon (Table 1) while the lowest temperature Marion Island, its value being more than twice that of Tristan da Cunha 105 Baardseth, ss 1941 recorded between 1951 and 1965 was 2, I ° C elements and offers no protection to any but the (Schultze,61 1971) According to Kno.x62 (1970) surface Gough Island. smallest forms of animal life. The oribatid mite waters immediately south of the Convergence have an The shores of Marion and Prince Edward are never Halozetes marionensis occupies micro-depressions in In spite of Marion's very limited range the rocky average temperature of about I-2°C. This means iced over though snow and ice pellets occur quite the Verrucaria thalli and though it may number up to shore zonation is well developed and follows closely that the Convergence must lie very close to Marion at frequently during winter. 12/ cm2 it is inconspicuous because it is small (800 p.m the pattern observed on other sub-Antarctic islands. times. in length) and dark grey. Occupying the same niche in The raising of effective tidal heights by wave action is Deacon's63 ( 1937) isohaline trace iri the vicinity of The littoral biota of Marion Island the lower half of H. marionensis' range another mite a well known phenomenon, and the contention of Marion indicates a salinity of 340/ between Transects (Mesostigmata) was encountered. Bright orange in 00 colour, it is more conspicuous than H. marionensis Stephenson & Stephenson59 ( 1949) that the tides do November 1931 and March 1933. Orren64 (1966) The transects are presented in order of increasing though as tiny. The highest concentrations of 18/ cm2 not directly cause zonation is well borne out by obtained salinities of 33,69°/00 at ten stations in the complexity from the evenly sloping unbroken shores observations on the west coast of Marion where swell vicinity of Marion and the Crozets for June/ July of Macaroni Bay to the shaded boulder-strewn cove occur in shallow grooves running parallel to the height consistently overrides the tidal range by 1962. Inshore salinities measured by Fuller65 ( 1967) at Duiker's Point. Figs. 3- 7 show the profiles of the shore. A few specimens of the siphonarid gastropod several metres. for the period January to March 1965 ranged from transect sites, and the commonest species are Kerguelenella lateralis inhabit larger depressions and 32,80/00 to 33,5° I 00 • sketched diagrammatically to illustrate the general Swell zonation. Relative abundance and detailed distribu­ grooves in the rock at the lowest level of the lichen zone. In summer the middle section of the lichen zone During recent years meteorology staff based at the H ydrology of tide pools tion of organisms are indicated by bands of varying is overlain by a continuous cover of a chlorophyte Marion weather station have estimated swell height, Tide pools of considerable size are infrequent on width in the upper halves of the figures. Zone­ Ulothrix sp. which appears bright green on drying period and direction daily in addition to taking sea Marion. At the southern end of Transvaal Cove a forming species typical of the sub-Antarctic regjon were used as indicators of effective tidal levels and no out during rare periods of sunshine. Table J attempt was made to relate these levels to chart Porphyra zone (upper littoral to mid-littoral): In datum. Reasons for such procedures have been addition to inhabiting the lichen zone the orange mite Average and extreme sea surface temperature (0 C) at Marion Island 1951 - 1965 explained by Lewis68 ( 1972) and have been employed (Mesostigmata) occurs throughout most of the in the sub-Antarctic by Knox,69 (1960), Kenny & Porphyra zone and reaches its lower limit shortly J F M A M J J A s 0 N D Year Haysom7o (1962) and others. The zonation of before Porphyra becomes scanty. Kerguelenella Macaroni Bay Transect I was regarded as the basic lateralis is more common but remains confined to Average 5,7 o.l 6,1 5,7 5,0 4,7 4.3 4,0 4,0 4,2 4,7 5.1 5,0 pattern on Marion and descriptions of other transects grooves and d epressions. At the lower limit of Maximum 6,9 7,9 8,0 7.7 6.4 6.4 6.3 5.1 5.7 5.4 5,9 6.5 8,0 are referred to in terms of departures from this basic Porphyra deep crevices harbour small numbers of the Minimum 4,4 4,3 4,6 4,0 3.0 3.1 2.5 2,6 2,1 2.~ 2,6 3,8 2,1 pattern. Discrepancies in tidal levels reached by pelecypod Kidderia bico/or. A few specimens of the 8 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 9

Kelp zone (lower tittoral): Very little overlap between organisms above and below Durvi!lea antorctica occurs and this large alga forms a natural break on the shore. (In this work Durvillea has been referred to the species antarctica but a much-needed revision of the genus on a world-wide basis may invalidate this.) Kergue!ene!la lateralis reaches its lower limit a few centimetres below the upper limit of the Durvil!ea holdfasts. Shells at this level are encrusted with algae while those of the lichen and Porphyra zones are bare indicating that Kergue!ene!!a does not migrate vertically but has a wide vertical range. Kidderia hicolor extends into the kelp zone in small numbers and remains confined to crevices and depressions. Nace!!a delesserri congregates in a narrow belt about 40 cm wide from the upper limit of Durvillea downwards. Amphipods are common and many take refuge in the pallial cavities of Nace!la. Branch75 ( 1975) in a study of South African limpets suggests that juvenile Hya!e grandicornis avoid desiccation at low tide in this way. On Marion this behaviour would be adaptive in eliminating osmotic stress during heavy rain. cm -- About 10 cm below the upper limit of Durvil!ea ---- lithothamnia commence and cover the substrate as a Enteromorpha• I sp. continuous encrustation down into the sublittoral. Scattered tubes of serpulid worms (Spirorbinae) occur occasionally on the lithothamnia and a few planarians were noted. Inconspicuous but fairly 481 ~ common is the tiny amphineuran Hemiarthrum setulosum which adheres to the surface of the lithothamnia but may extend higher to near the upper Plate 2. Macaroni Bay Transect I. Porphyra and Durvillea limit of the holdfasts of Durvillea. amarc1ica are well developed ..

3M ~ Durvi!lea antarctica is very well developed and a patellid Nace/la delesserti extend marginally higher count of 94 mature plants was obtained along I 0 m of than the upper limit of the Dun1i/lea anrarctica this shore (5 m on either side of the transect line). It is holdfasts. N a cella thus represents the first non­ necessary to cut several plants away before observa­ cryptic macrofauna encountered. Amphipods are tions and collections can be made since the fronds 2M ~ abundant in the mid and lower Porphyra zone. Near cover the rock as a dense and continuous blanket. the lower limit a sample of Porphyra was scraped Organisms beneath the fronds receive a measure of from 400 cm2 (20x20) of the rock and scrutinized for protection from the force of the swell and the 202 ~ fauna. A count of 96 amphipods. principally Hya!e elements, but live under conditions of heavy shade. grandicornis and H. hirtipa!ma, was obtained. In the During rough seas the fronds whip back and forth mid-Porphyra zone they numbered 69/ 400cm2 while and abrade the substrate to at least HWN. The fronds numbers had declined to 9/ 400cm2 near the upper and stipes are devoid of macroscopic forms of life but limit of Porphyra. the holdfasts of older specimens may harbour 1001 ~ The encrusting Verrucaria of the lichen zone above molluscs. crustaceans and polychaets. extends into the Porphyra zone but becomes patchy Lower red zone (Upper sub-littoral): It was not and is taken over towards the lower limit of Porphyra possible to work efficiently below Durvil/ea on this by encrusting growths similar in appearance to shore and the level labelled -58 cm on the transect Verrucaria. These were not distinguished in the field Of ~ diagram was estimated visually and not visited. so it is not yet possible to accurately define the lower Between swells glimpses of the lower red zone limit of Verrucaria on the shore. At least two species revealed scattered specimens of Desmarestia chor­ -58 r:;------~ of lichen V. microspora and Arthopyrenia sublit­ da/is and D. rossii below a continuous carpet of 753 cm tora!is extend to the Durvil!ea zone on the Auckland mixed red algae. notably Schizoseris sp. islands (Knox, 74 I 968), where algae such as Abroteia Prasiola crispa suborbicu!aris may occur as well. Abroteia, inter­ ~ Porphyra sp. Abundant ~ spersed with some Verrucaria, was in fact recorded MACARONI BAY TRANSECT 2 ~ Verrucaria spp. ~ Enteromorpha sp. from this level elsewhere on Marion. Further detailed Here the grey lava slopes gently seaward. There is no • Common collections of these important encrustations are major departure from Transect I at Macaroni Bay as required to resolve their components accurately. ~ Lithothamnia - Fairly common far as general zonation is concerned though addi­ The extensive band of Ulothrix also extends well tional species appear, sometimes in considerable -Rare ~Schizoseris sp. into the Porphyra zone, and scattered growths of abundance (Fig. 4; Plate 3). The most obvious minor Enteromorpha bu!bosa occur at the lower limit of differences are: Fig. 3. Ma<.-aroni Bay Transect I. The distribution and relative abundance of the major zone-forming organisms. Porphyra. 1. The blackening encrustation of lichens (and 10 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 11

possibly other organisms) is conspicuous from the 2. Mites (Acarina) are not common on this shore. lower supra-littoral to near the upper limit of 3. Rhodymenia species I is fairly common towards Durvillea antartica because Porphyra is poorly the lower limit of Porphyra though it does not developed. form a distinct band as described for Macquarie

K erque/enella

Lasaea consanguinea Plate 3. Macaroni Bay Transect 2. Note shallow pools, sparse Porphyra but well developed Durvilfea antarcrica.

Island by Kenny & Haysom16 (1962). (Two in shallow depressions. A few small pools in the mid Rhodymenia species occur in the upper and mid­ Porphyra zone contain dense growths of Entero­ littoral of the island. Species J is luxuriant and morpha bulbosa and some Phyllophora appendicu­ frondose; Species 2-see Duiker's Point Transect lara. Amphipods (Hyale spp.), isopods and the - short-cropped and turf-like). copepod Tigriopus angulatus are common in the 4. Luxuriant growths of Leptosomia ?simplex cover pools. Lasaea consanguinea inhabits the filaments of the rock between Durvillea holdfasts from the Rama antarctica which occurs in crevices. Also middle to the lower horizons of the Durvillea associated with crevices and depressions but not with zone. Rama are Kerguelenella lateralis, Kidderia bicolor and occasionally serpulids (Spirorbinae). Amphipods Details of the zones (Hyale sp.) are common in the lower region of the Lichen zone (lower supra-littoral): Lichens, principal­ Porphyra zone. Near the lower limit of Porphyra ly of the genus Verrucaria, occur as a continuous pools allow the extension upwards of organisms from black encrustation. Scattered growths of a poorly lower levels and the amphineuran Hemiarthrum developed light brown form of Porphyra may be setulosum was found together with juvenile patellids found in the lower half of the lichen zone while Nacella deiesserli. Ulothrix sp. is abundant at this level also. Shallow Kelp zone (lower littoral): The gradual slope of this crevices harbour dense tangles of Rama antarctica. stable shore allows optimum development of The filaments of this chlorophyte are colonized by Durvillea antarctica and the great fronds lie in numerous specimens of the pelecypod Lasaea profusion on the rockface. Many plants have to be consanguinea and to a lesser degree by the cut away before effective observation or collecting oligochaet Lumbricillus aestuum. The siphonarid can be carried out. A dark brown encrustation Kerguelenella lateralis extends into the lichen zone elsewhere identified on Marion as comprising the from lower levels and occurs in cracks and alga A broteia suborbicularis, an unidentified alga, depressions whi'ch may also contain a few pelecypods, and the lichen Verrucaria mucosa, surrounds the notably Kidderia bicolor. holdfasts of the uppermost Durvillea. Lithothamnia Porphyra zone (upper littoral - mid-littoral): adhere to the substrate as a continuous sheet from Difficulty was experienced in defining the upper limit near Durvillea's upper limit to the sub-littoral. A of Porphyra with precision since unlike on the steep conspicuous alga forming a clearly defined belt in the shore of Transect 1, its abundance here diminishes mid Durvillea zone is Leptosomia ?simplex. It does gradually in the region of HWS. A stunted light not extend to L WS. brown form extends into the lichen zone, but was Several epiphytes are common on the stipes of 1040 cm disregarded since it did not conform to the Durvillea, notably Porphyropsis vexillaris, Ectocar­ appearance of typical Porphyra characteristic of pus sp. and yet another Rhodymenia species. Outliers Rhodymenia sp. HWS elsewhere on Marion. from deeper levels find their upper limit in crevices Porphyra is relatively poorly developed on this and conditions of shelter in the Durvillea zone, e.g. Verrucaria spp. tosomia sp. • Common shore and interspersed throughout by blackening Schizoseris sp., Lophurella sp. and Rhodoglossum Lithothamnia Fairly common Rare encrustation and Vlothrix sp. In the lower region of sp. Specimens tentatively identified as belonging to the Porphyra zone Rhodymenia species I is quite the genus Delesseria were also encountered. Fig. 4. Macaroni Bay Transect 2. The distribution and relative abundance of the major zone-forming organisms. common and grows luxuriantly on level surfaces and The abrasive action of the fronds of Durvillea 13 12 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 precludes the development of all but the hardiest only conspicuous animals in this zone. Crevices fauna on exposed surfaces, and Hemiarthrum however harbour a rich fauna, particularly the setulosum and juvenile Nacella defesserti are the pelecypod Kidderia bicolor and to a lesser degree K.

Plare 4. Ship's Cove Transecr. Porphyra is sparse and Durvillea amarcrica absent round the sides of landward boulders.

oblonga. These tiny organisms may fill a crevice to a lateralis on the seaward face of boulder 4. depth of several centimetres. Alse present are 2. Luxuriant growths of Porphyra and an associa­ serpulids (Spirorbinae), amphipods and isopods. The tion of Polysiphonia, Schizoseris and Myriogram­ Porphyra sp. asteroid Anasterias rupicola is rare and inhabits me commencing on boulder 3, with the disappear­ depressions towards the lower limit of Durvil/ea ance of Bostrychia vaga. where outliers of sub-littoral organisms such as 3. The appearance of Durvillea antarctica midway sponges, bryozoans and actiniarians may also be round boulder 2. Prasiola crispa found. Zonation at boulder I is the same as that of Macaroni Bay transect I , but the differences Lower red zone (upper sub-littoral); Delicate noted above occur on boulders to landward. rhodophytes such as Lophurella, Schizoseris and Myriogramme cover the substrate from Durvillea's Details of the zones lower limit at L WS down into the sub-littoral. Lichen zone (supra-littoral): The upper supra-littoral Scattered specimens of Desmarestia rossii attain their is characterized by orange, grey and white lichens as upper limit at L WS also. well as scattered growths of Prasiola crispa. Verrucaria spp. occur in abundance throughout the SHIPS COVE TRANSECT lower supra-littoral on boulders 5 and 4. Here too The transect site of Ship's Cove, (Fig. 5; Plate 4) isolated tufts of very poorly developed Porphyra may situated on a north-facing shore about 2,25 km north­ be found. The siphonarid Kerguelenella lateralis west of Transvaal Cove, was composed of large attains maximum abundance near the lower limit of boulders (up to 280 cm diameter) lying fairly closely Verrucaria and does not extend landwards beyond packed on a gently sloping substrate. A shore of this the seaward third of boulder 4. The oribatid type affords useful conditions for the study of wave Halozetes marionensis is not common on this shore. action since mean unbroken sea level (as opposed to Verrucaria is conspicuous on the summits of boulders mean wetted level) extends from the seaward boulder 3 and 2 where it is overlain in part by a dense growth landwards around the other boulders of the transect. of Ulothrix sp. Each boulder (except number 5) is thus subjected to Porphyra zone (upper littoral): Porphyra grows the full range of astronomical tidal fluctuations while luxuriantly on boulders 3 and 2 but somewhat sparsely on boulder I . Towards the upper limit of its -::;;;:::--- Grey lichen ~~~~~~~' Bostrychia vaga »i'1r' Ulothrix sp. the more seaward boulders become progressively influenced by the addition of wave action. Surfaces of range it is interspersed with abundant Ulothrix sp. ~ Prasiola crispa :.:::::. Abroteia I Verrucaria .. Kerguelenella lateralis boulders closely packed together could not be On the seaward face of boulder I, light growths of examined efficiently, but zonation on such surfaces Enteromorpha bulbosa compete with Porphyra and I2J2ll1J2 Porphyra sp ~~\~\') Lithothamnia .!.!__ Nacella delesserti was influenced by increased shade as well as wave extend to the Durvillea holdfasts. The orange mite ·:.:·::.::.·.· Verrucaria spp. action. (Mesostigmata) is fairly common to leeward of ~\.\ Pol~siphonia, Schizoseris .W~ l\~ Enteromorpha bulbosa _If_ Mynogramme ' The pattern of zonation from the upper littoral to boulder I near the summit and extends landwards on the upper sub-littoral undergoes three major changes boulders 2 and 3 from their summits to near the lower • Common limt of Porphyra. • Abundant - Fairly common - Rare between the most landward boulder and the most seaward boulder of the transect. These changes may Bostrychia vaga zone (mid-littoral - lower littoral): be summarized as follows: Bostrychia vaga is confined to the sheltered conditions at boulders 5 and 4 and occurs as a carpet Fig. 5. Ship's Cove Transect. The distribution and relative abundance of the major zone-forming organisms. I. The appearance of abundant Kerguelenella 14 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 15 on slopes that face the substrate at various angles. common. Careful searching among these boulders continuous encrustation where a few fronds of very grandicornis and H. hirtipa/ma does occur, H. Against the substrate beneath the B. vaga on boulder revealed in parts aggregations of serpulids Helicosi­ poorly developed Porphyra may be found as well.At hirtipalma is the commoner species lower on the 5 a few oligochaets and insect pupae occur, while on phon sp. and a few Paralaeospira levinseni which had the upper limit of Porphyra, V. tesselatula thins and shore particularly among the fronds of Rhodymenia boulder 4 the filaments of the alga harbour large on the larger tubes of Helicosiphon. A few soon disappears. species 1. Here gross counts of amphipods yielded numbers of the pelecypod Lasaea consanguinea as actiniarians were noted between the small boulders, The homogenous appearance of the black lichens 392/400 cm2. well as oligochaets, Lumbricillus sp. and the tiny fish Harpagifer bispinis is not underlies what may be a very complex zone A few specimens of ?Allorchestes sp. were noted Only the seaward and leeward faces of boulder 4 uncommon. composed of several overlapping and intermingling among the Rhodymenia fronds as well as the tiny could be examined at this level since the other The raising of effective tidal height by wave action species. More detailed sampling from this area is oligochaet Lumbricillus aestuum. surfaces were too close to surrounding boulders. is well illustrated at the seaward end of the transect needed. Kelp zone (lower littoral): Durvillea antarctica is Such surfaces have been left blank in the diagram where an upward trend of zone-forming organisms During rare periods of sunshine the lower half of not abundant on this shore as a result of instability of (see also boulders 5, 2 and 1). Between the lower limit may be noted. the lower lichen zone appears bright green due to the substrate. Where it does occur it favours larger of Verrucaria and the upper limit of Bostrychia on extensive coverage of the boulders by the chlorophyte boulders or those firmly wedged between other the seaward face of boulder 4, the brown encrusting TRANSVAAL COVE TRANSECT Ulothrix sp. boulders. Just below Durvillea's upper limit litho­ Abroteia/ Verrucaria may be found. The transect site at Transvaal Cove (Fig. 6; Plate 5) Common organisms of the lower lichen zone are thamnia coat the rocks and extend down into the sub­ Abroteia/ Verrucaria zone (lower littoral): The situated below the meteorological station on an east the oribatid Halozetes marionensis and the orange littoral. Extensive bare patches indicate the frequency calm conditions that prevail at boulder 3 allow facing shore, was composed of boulders ranging mite (Mesostigmata). H. marionensis occupies of rotation and abrasion. An alga commonly noted in optimum development of the Abroteiaf Verrucaria between 50 and 70 cm diameter. The boulders lie depressions in the Verrucaria tesselatula thalli and the Durvillea zone is Leptosomia sp. but it does not complex where it extends as a sheet from the upper piled on top of one another along a stretch of I 00 m occurs in dense aggregations numbering up to 141 cm2 reach the sub-littoral. limit of lithothamnia to near the level reached by forming a boulder beach. at the middle of its range. It extends up the shore with Juvenile specimens of the patellid Nacella delesserti Kerguelenella lateralis. The zone becomes progres­ Lack of light and severe restriction of growing V. tesselatula but does not reach higher than the are common throughout this zone and attain peak sively narrower to seaward and at boulder I is all but space preclude the ·development of algae down the upper limit of this lichen where its numbers drop to numbers a little below Durvillea's upper limit. A obscured by the holdfasts of Durvillea antarctica. sides of the boulders as at Ship's Cove. Swells of less than 1I cm2. At the lower limit of its range H. surprising feature of this shore is the absence of Tolerant of shady conditions, Abroreia/ Verrucaria above average strength rotate the boulders so that marionensis is confined to the landward sides of Hemiarthrum setulosum. An extensive search did not 2 was noted on stones between boulders 5, 4 and 3. abrasion further limits growth. A degree of stability is boulders where counts of 2/ cm were obtained. yield a single specimen. According to Simpson77 Apart from a few Nacella delesserti no organisms achieved against the substrate beneath several layers Orange mites compete with H. marionensis on the (1969), Hemiarthrum shows a strong food preference were found to occur on this encrustation. of boulders, and on stones which have fallen between Verrucaria in the lower half of H. marionensis' range for coralline algae. The patchy distribution of Kelp zone (lower littoral): The calm conditions at the boulders and come to rest on the substrate. Here a and number 1-21cm2• Further down the shore they lithothamnia on this transect may thus be limiting. A boulders 5 and 4 preclude the development of any variety of invertebrates may be found, but they are are commoner and reach peak numbers of up to few Kerguelene!la lateralis were found in depressions Durvillea antarctica and it is first encountered very difficult to reach and collect systematically. 21 f cm2 on the third boulder above the upper limit of near the upper limit of Durvillea. The surface of the midway round boulder 2 where a few holdfasts Zonation on the superficial layer of boulders is Porphyra. They extend a short distance into the lithothamnia-covered rocks is devoid of other forms adhere to the rock just above the level reached by similar to that of Macaroni Bay Transect 1 except Porphyra zone. of macroscopic life. encrusting lithothamnia. To seaward Durvil/ea that Rhodymenia species I forms a belt between The siphonarid Kerguelenella lateralis is fairly The boulders of this transect continue into the sub­ becomes more common and attains maximum Porphyra and Durvillea. common in the lower half of the zone but it inhabits littoral but initially the slope is far more gradual than density on the seaward face of boulder I where two On this shore the distribution of organisms was pock marks and depressions in the boulders and is in the littoral. Seventeen metres seaward from the holdfasts may occur one above the other. Nacella studied from the upper supra-littoral to the sub­ therefore most inconspicuous. upper limit of the lower red algae the depth was delesserti is a common organism found throughout littoral at a depth of 3,5 m. Porphyra zone (upper and mid-littoral): Porphyra found to be only 120 cm at low spring tide. the Durvillea zone, and its vertical range increases sp. is well developed on this shore and grows Turbulence and frequent rotation of boulders results with that of Durvillea on the seaward face of boulder Details of the zones luxuriantly on the more stable boulders. Towards its in distribution patterns of specially adapted organisms I, where a preponderance of juvenile specimens Upper lichen zone (upper supra-littoral): The upper lower limit it competes with Rhodymenia species I. so that the upper sub-littoral has a characteristic occur. Nacella extends landwards to boulder 3 and supra-littoral is characterized by a colourful mosaic Verrucaria tesselatula is hardly evident below the appearance. forms a clear line along the transition between of lichens comprising seven species. The most upper limit of Porphyra, but Ulothrix sp. extends The upper sub-littoral of Knox78 (1968) includes Abroteia/ Verrucaria and lithothamnia. During rough conspicuous of these is the bright yellow Caloplaca patchily through most of the Porphyra zone Macrocystis which on Marion does not generally weather Nacella may migrate upwards to near the cirrochrooides which extends from the lower limit of diminishing rapidly however below Porphyra's upper grow in less than 10 metres of water. Since at least lower limit of Porphyra. The amphineuran Hemiar­ maritime vascular plants to near the upper limit of limit. A few growths of Enteromorpha bulbosa occur some zonation occurs within this range the author thrum setulosum is common on lithothamnia the lower supra-littoral where it becomes confined to towards the lower limit of Porphyra, and just above has (quite arbitrarily) divided the upper sub-littoral throughout the Durvillea zone but ends abruptly with the landward sides of the boulders. Common the upper limit of the holdfasts of Durvillea into a Lower Red zone I and a Lower Red zone 11. Durvillea midway round boulder 2. throughout the range of Caloplaca is a white lichen antarctica a smooth dark brown encrustation coats Lower red zone I (upper sub-littoral): Durvillea Lower red zone (upper sub-littoral): The upper Pertusaria sp. Other lichens characteristic of this zone the rocks. This was found to contain two algae (one antarctica is largely absent, scattered immature plants limits of lithothamnia and Po(vsiphonia sp. coincide but which do no extend as far down the shore, of them unidentified), Abroteia suborbicularis, and occurring near LWS only. The cover of lithothamnia almost exactly at L WS on boulder 3, but on boulder include Lecidea interrupta and more rarely Buellia small quantities of Verrucaria mucosa. is complete on the upper surface and round the sides 2 and further seawards the lithothamnia reach far toniops and Haematomma erythromma. Occupying The orange mite extends marginally into the of boulders which have remained in the same position higher levels than Po(rsiphonia sp. The calm the level which marks the transition from the upper Porphyra zone where its numbers drop to 0-1 1cm 2 for a reasonable length of time. Recently rotated conditions to leeward of boulder 2 allow collections lichen zone to the lower lichen zone is a distinct but shortly before disappearing altogether. Kerguelenella boulders may be partially or almost entirely bare to be made in the upper sub-littoral where three narrow band of Verrucaria durietzii. the thalli of lateralis is fairly common throughout the Porphyra depending on the degree of rotation and the extent of distinct belts of rhodophyte species would seem to which are interspersed with small quantities of zone but the scattered nature of its occurrence the surface area which formerly faced the substrate. justify the designation of a sub-littoral fringe as a Verrucaria mucosa. At this level too the alga Prasiola precludes expressing it in numbers. Red algae are abundant throughout the zone, the definite zone. The three belts from L WS downwards crispa may be found. It favours rocky surfaces below Amphipods are very common among the Porphyra most conspicuous being a broad leafy Rhodoglossum are composed principally of Polysiphonia sp.. bird perches and extends landwards for a short fronds, the dominant species being Hyale grandicor­ sp. whose internal vegetative structure was found to Schizoseris ?dichotoma and Myriogramme sp. distance into the zone occupied by Caloplaca. nis and H. hirtipalma. Counts were obtained by be similar to Iridaea sp. Rhodoglossum favours the Present with the Poll'siphonia are small quantities of Lower lichen zone (lower supra-littoral): The lower scraping measured areas (20 x 20 cm) of Porphyra tops of boulders. Round the sides and in places Ballia callitricha, Lophure/la sp. and Rhodog/ossum lichen zone is easily recognizable from afar as a broad and then removing the amphipods individually once partially hidden by Rhodoglossum may be found sp. These separate belts are not illustrated on the black band forming a continuous covering from the preserved. H. grandicornis numbered 6371400 cm2 at Polysiphonia urbanoides, Lophurel/a sp., Myrio­ diagram. Smaller boulders against the su bstrate Porphyra up the shore to within a few metres of the the middle of its range in the mid Porphyra zone. A gramme sp. and Schizoseris sp. Rarer algae of this between the major transect boulders are lean in lichens of the upper supra-littoral. The principal preponderance of juveniles occurs near the upper zone include ,Spongomorpha pacifica, Ceramium branching algae but are well covered by Iithothamnia. component is Verrucaria tesselatula. Scattered thalli limit of Porphyra where numbers declined to rubrum and Geminocarpus geminatus. To leeward of boulder 3 Codium sp. becomes quite of V. durietzii extend to near the upper limit of the 432/ 400 cm2. Though some overlap between H. Living among the fronds of these algae are large S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 16 S. Afr. J. Antarct. Res., 1976, Suppl. 1 17 numbers of amphipods, principally Hyale hirtipalma boulders as well. Here Exosphaeroma gigas is quite and Jassa fa/cat a. Round the sides of the boulders on common and favours the substrate under the lithothamnia and sponges, Pontogenia sp. was boulders. Juvenile Nacella delesserti are present at commonly noted. The isopod Dynamenella hurtoni this level where they adhere to the ljthothamnia. also lives in close association with the algae but may Many littorinids Laevilitorina caliginosa and more extend to the substrate and interstices between the rarely the pelecypod Gaimardia trapesina were taken

c 0 E E c 0 0 () E > E ~ G) 0 · ~ i .,:::. - l _'ll = - - - :: I - I ' I I I '<. (.) U. a: ~ Pla1e 5. Transvaal Co,·e Transec1. Nole scauered Porphyra and Dun·i/lea amarctica. Boulders cast up from 1he sub­ -~---·o:: .() ... lit! oral bear patches of bleached lilholhamnia. E ~ :::. ~

-J ~ •• , ·~; ~ ~ .; j I I l ~ ! /: ~ E I ·;:"' II ·;;:"' '"'E from among the fronds of the algae. along a horizontal distance of 56 m from the L WS I "' E ~ 0 It was possible at this level to investigate the level on the shore. Two major changes in zonation I E:... .s::"' CO I c I Q o undersurfa.ces of the superficial layer of boulders and may be noted. The first occurs shortly below about I "J:: £- ·e I Cl: :J ... the interstices between the boulders. Common 100 cm below L WS; the second in the region of the I ~ organisms noted were sponges, particularly the deepest point (300-350 cm below L WS) some 50 m f "c I mauve Haliclona topsenti which had tiny yellow from the shore. I ...~ I 0 isopods living on it. Other common sponges include Below I 00 cm below L WS the typical algae of the I "(? Clathrina coriacea and Leuconia joubini. The lower red zone I all but disappear, their place being 8 pycnogonid Tanystylum cavidorsum creeps over largely taken over by yet another Polysiphonia I u .r:. lI lME " encrusting growths and was also taken from substrate species, Geminocarpus geminatus, Ballia callitricha '- a. ~ • I I I ~ 0 detritus. Delicate hydroids seek the shelter of and Ceramium rubrum. The flat encrusting lithotham­ "' -- ~ I .~ I I I~ ~ "() interstices and undersurfaces of boulders, the most nia characteristic of the littoral give way to the more I ..c "0 luxuriant Phymatolithon sp. which builds up as c: conspicuous of these being Myriothela meridiana. ~0 ' I iI lI Iij j j.. :3 I u .0 The tiny Sertulare/la picta grows in abundance from concentric overlapping ridges. Durvillea antarctica is ~ I ' I I "0 e os the sides of boulders. Further hyroids commonly altogether absent and the sponges disappear. 1l . ~ noted belonged to the families Tubulariidae and With increasing depth the substrate stabilizes and X ; ~ ~ Hydractiniidae, but lack of mature gonophores encrusting growths of corallines, lithothamnia and precluded identification. Actiniarians are quite other algae spread between and over successive "c .. common under boulders and a single beige nudi­ boulders to form a uniting matrix. c t .2 branch was found . In the region of the deepest point of the transect :; Echinoderms are poorly represented at this level, (300-350 cm) very large algae appear in abundance, I ~ i\ l ! i lll .0 :§ '5 only the holothurian Pseudocnus laevigarus and a particularly Durvillea ?antarctica, Desmaresria rossii . ~ " single specimen of the asteroid Anasterias rupico/a and D. chorda/is. On smaller boulders between these ... being recorded. Polychaet worms attain high algae adult Nacella delesserti congregate in huge ~ 1;) ~ 'i5 ·~ i i I i ~ numbers on the sides of the boulders and among numbers one on top of the other to form extensive .. a·§ I ;:; e ~ I I I I I 2 "t; :::. I I I I J ~ detritus underneath. The most conspicuous of these is carpets several metres in extent. The cover of ""'c "' o t: t 1 I I ., "' the tube-building Platynereis australis and the scale lithothamnia is largely obscured by Corallina sp. ci • .% .~ ~ i): I I I I ~ · ~ r-e "' t u I n( I I I I I ·s ... worm Harmothoe bergstroemi. Sifting of grit from which favours the larger stable boulders. Other algae ~ ~ beneath the boulders yielded Nicolea chilensis, collected from this area include Ulva sp., Ballia ..,S! 8 Eulalia magalhaensis, Cirratulus cirratus, Platy nereis cal/itricha, ?lridaea, Schizoseris sp., Halopteris ~ .. ~ australis and Syllis sp. In this niche Nemertea funicularis, Ceramium rubrum; Rhodophyllis sp., ~ including Tetrastemma sp. were found to be fairly PolysijJhonia sp. and Heterosiphonia sp. A dive to a ~ common and a few sipunculids Golfingia ohlini were depth of 8 m (beyond the transect line) revealed >t:i also encountered. extensive coverage of the boulders by coralline algae eO Lower red zone ll (upper sub-littoral): Collections and scattered large brown algae. The levels ii: were carried out to a depth of 350 cm at 8-m intervals investigated were probably insufficient to show up 18 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 19 any major depth zonational changes and those and Kergue/enel/a are encountered at shallow lower on the shore the orange mite (Mesostigmata) limit of Dun·illea and the upper limit of the lower red recorded are probably correlated with size of depths on the under-surfaces of the boulders while may be found in small numbers. A surprising feature algae. Ballia callitricha is the principal constituent of boulders and substrate stability. A general descrip­ serpulids, Kidderia and Nacel/a follow in succes­ was the discovery of numerous tiny pelecypods the lower reds. Its fronds may be very heavily clothed tion of the lower red zone II follows. sion below. Exosphaeroma inhabits the substrate Lasaea consanguinea and a few Kidderia bicolor with a blue green alga and occasional tufts of Amphipods are common throughout among the beneath the boulders. among the tangled fronds of Bostrychia and Rama: epiphytic Heterosiphonia sp. Ectocarpus sp. is fronds of red algae, both Hyale hirtipalma and Jassa 3. Above Verrucaria zone (upper supra-littoral): one would not expect to find filter-feeding organisms common as smooth rust-coloured mats at this level falcata being recorded to 350 cm. The isopod Encrusting lichens occur patchily for 30 cm so high on the shore. Oligochaets Lumbricillus sp. and Plocamium sp. occurs occasionally among the Dynamenella huttoni was noted commonly here also. followed by a scattered green stain-like adherence and insect pupae were also found in this niche. The Ballia together with Delesseria sp. The fronds of algae also harbour large numbers of the for approximately 20 cm where a fe'-' pelecypods siphonarid Kerguelenella lateralis occurs in small Four species of sponge occur between and around littorinids Lavilitorina caliginosa and Eatoniella Lasaea sp. were noted. At 60 cm depth numbers throughout the lichen zone. the sides of boulders. They are Leucosolenia kerguelensis and more rarely the pelecypod Lissarca oligochaets. planarians, amphipods and a few Porphyra zone (upper and mid littoral): Though hotryoides, Amphilectus fucorum, Ectyomyxilla chi­ rubrofusca. isopods occur and these persist to the level of grit Porphyra is very poorly represented on this shore the lensis and Halichondria panicea. The amphipod The great number of Nacel/a delesserti at the deep against the substrate. name will be retained for the sake of uniformity. Jassa falcata lives among them in considerable limit of the transect has been referred to, but they Encrustations of blackening organisms such as numbers while algae at this level are colonized by occur less commonly in shallower depths also, some numerous Pontogenia sp. The isopod Exosphaeroma DUIKER'S POINT TRANSECT Verrucaria occur throughout this zone reaching to favouring the upper surfaces of boulders, others the upper limit of Durvillea antarctica holdfasts. gigas occurs under stones. Harpacticoid copepods occurring round the sides where the rocky surface is The Duiker's Point transect (Fig 7; Plate 6) was Compact masses of several species of algae growing live in profusion among the fronds of Ballia. situated just over a kilometre north of Transvaal devoid of encrusting growths. The shells of Nacella in close association with one another from turf-like /femiarthrum sewlo.H/111 is common on the lithotham­ on top of boulders are covered with lithothamnia Cove in a small cove some ten metres wide and eight carpets on top of the boulders. The principal metres deep. The cove is floored by boulders of nia together with Nacel/a delesserti. Crevices and while those on the sides of the boulders have clean components are Rhodymenia species 2, Cladophora varying sizes, many of them only 30-40 cm in dissections among the boulders harbour abundant shells suggesting that this organism does not migrate sp. and Rama sp. Further down the shore diameter, which facilitate removal and collection at Kidderia bicolor which may also form aggregations vertically. Juvenile Nacella may be noted throughout Rhodoglossum sp. appears among the turf and deeper levels. Steep cliffs on either side of the cove cut against the substrate under encrusting sponges. the sub-littoral transect but become very rare near the becomes dominant near the upper limit of Dun•illea. down light considerably. Fresh-water seepage occurs Occasional specimens of Kerguelenella lateralis were deep limit of the transect. At this level Rhodymenia species 2 and Cladophora down the cliff faces. The cove itself leads off from the noted in this zone. Kerguelenella thus has an The asteroid Anasterias rupicola is fairly common disappear but small quantities of Rhodymenia species exceptionally wide range. Echinoderms are not and frequently preys on Nace/la. Pseudocnus landward end of a curved alley about 30 m long and 8 m wide. The seaward end of the alley has direct I may be noted in isolated patches. Other algae common on this shore, Anasterias rupicola and laevigatus occurs under boulders. Around the side of associated with the turf in small quantities include Pseudocnus laevigatus being recorded only in small boulders under protected conditions bryozoans were communication with the open ocean and faces east. Luxuriant development of Durvillea antarctica at Enteromorpha bulbosa. Bostrychia mixta and Ecto­ numbers. The hydroids Myriothela meridiana and encountered and provisionally identified as Menipea carpus sp. The density of the turf is further increased Silicularia rosea occur around the sides of boulders in crispa. Hydroids may be noted in this niche also. the transect indicates persistently strong wave action. However the cove is protected from violent storms so by an accumulation of algal fragments of Ballia shelter where actiniarians are quite common as well. Various polychaets including Harmothoe berg­ callirricha, Sphacelaria sp. and Polysiphonia sp. cast Polychaets are well represented at this transect and stroemi and nemerteans occur under boulders. that boulders in the Dun·illea zone are not frequently rotated. up from deeper levels as drift. Single thalli of are nearly all confined to the undersurfaces of Fishes are quite common, particularly Notothenia Porphyra occur here and there. boulders and the substrate. The commonest include coriiceps which lies motionless between boulders and Zonation on the superficial boulders of this transect differs from the basic pattern in the following The orange mite (Mesostigmata) extends a short Romanchella inventi.~. Platynereis australis, Syllis may be touched by hand. Harpagifer bispinis lives distance into the Porphyra zone from the lichen zone, respects: variegata. Po~rcirrus hamiltoni. Cirratulus cirratus under stones while a few specimens of Notothenia but Kerguelenella lateralis persists right to the and Boccardia polrhranchia. The serpulid Roman­ macrocephala were encountered in mid water. 1. In the lower lichen zone dense tangles of Durvillea holdfasts and to deeper levels. Tiny chella was also found in aggregations underneath Further sub-littoral collections at other points and Bostrychia spp. and Rama sp. occur with oligochaets Lumbricillus macquariensis and L. encrusting sponges. A single hirudinean G(Jptono­ examination of holdfasts of Macrocystis and Verrucaria. aestuum inhabit the algal turf mat where a few tobdella antarctica was collected from under a Durvillea cast up after storms have brought 2. Porphyra is virtually absent, its level on the shore amphipods occur as well. The pelecypods Lasaea boulder. organisms such as hirudineans and ascidians to light, being taken over by a turf-like mixture of consanguinea and Kidderia bicolor occur among the In-fauna above L WS: Conditions between the and the shortcomings of snorkel diving for specimens Rhodymenia species 2, Cladophora sp. and Rama algae to near the upper limit of Dun•illea. boulders and on the substrate become progressively on a transect are fully recognized. sp. Depressions and grooves contain small numbers of calmer towards the landward cliff face as the force of 3. The lower lichen and the P01phyra zones occupy Kidderia sp. which may also be found in the upper the swell is dissipated. Natural sorting of the boulders Vertical zonation on the shore largely the same level in the vertical plane but are sub-littoral under sponges. during violent storms results in a vertical gradation of The difficulty of removing several layers of large separated on the horizontal plane. Kelp zone (lower littoral): The upper limit of particle size with the smallest particles accumulating boulders to reach organisms at deeper levels vertically 4. In the lower red zone Ballia callitricha is the Dun•illea antarctica is clearly defined midway down against the substrate. At this level there is a build-up below algal and lichen communities has been referred dominant alga. the large seaward boulder of the transect. Around the of organic detritus and crushed shells which retain to. This was however achieved in part at the upper holdfasts may be noted the brown Abroteia/ Verruca­ water at low spring tide. This together with the limit of Porphyra, in the mid Verrucaria 1one and a ria encrustation against the rock face. It is soon relative stability of the boulders allows optimum short distance above the Verrucaria zone. Details of the zones replaced by a continuous cover of lithothamnia. development of interstitial and substrate organisms. I. Porphyra zone: About half way round the sides of Lower lichen =one (lower supra-littoral): Verrucaria Hemiarrhum sewloswn lives mainly on the surface Lithothamnia coat the boulders where even small boulders, the Verrucaria disappears leaving the sp. coats the boulders sporadically in the upper of the lithothamnia. a few specimens penetrating amounts of light penetrate and extend from the rock faces devoid of encrusting forms of life. reaches of the lower supra-littoral but becomes higher to near the upper limit of Durvillea. Juvenile seaward boulder to near the cliff face. Nacella Kerguelenella lateralis occurs in small numbers, commoner near the upper limit of algal turf. Against Nacel/a delesserti occur commonly alongside He­ delesserti is common around the sides of boulders. and at a depth of 50-70 cm a few amphipods the cliff face and on the tops of boulders rich growths miarthrum throughout its range. Hemiarthrum may Where they occur on lithothamnia-free surfaces their Hyale grandicornis and serpulid worms (Spiror­ of Bosrrychia vaga mixed with nearly as much B. also be found under the holdfasts of mature shells are also free from lithothamnia, otherwise they binae) were noted. Between 70 and I 00 cm the mixta and B. typica occur. Rama antarctica and Durvillea. Two polychaets Cirratulus cirratus and bear dense encrustations, suggesting they move little. boulders become far smaller (e.g. less than IS cm Enteromorpha bulbosa colonize crevices and depres­ Boccardia polybranchia were taken from grit that had The under-surfaces of stones and small boulders diameter) and are colonized by dense aggrega­ sions at the same level. Rama antarcrica may grow accumulated under the holdfasts. A rich in-fauna against the substrate bear numerous planarians. An tions of small pelecypods Kidderia sp. and epiphytically on Bostrychia vaga and these two algae occurs between the boulders and among detritus amphipod Pontogenia sp. is common against the abundant serpulids. At levels deeper than 100 cm extend a metre or more up the cliff face where a thin against the substrate at levels just above LWS. A substrate and among stones. The isopod Exosphae­ adult Nacella delesserti are fairly common but coating of what was provisionally identified as description of this fauna is given later. roma lives under boulders throughout this zone and juveniles rare. The isopod Exosphaeroma gigas Acrochaetium and Rhodochorton may be found. Lower red zone (upper sub-littoral): Lithothamnia is particularly common near the landward cliff under attains high numbers against the substrate. Halozetes marionensis lives on boulders encrusted coat the boulders throughout the lower red zone and piles of decomposing Durvillea antarctica. The 2. Verrucaria zone: As in the Porphyra zone Hyale with Verrucaria, but is not very common. Slightly are the only growths on the rocks between the lower commensal isopod /ais pubescens is abundant on 20 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 21

Exosphaeroma. Harmothoe bergstroemi. Two species of amphineuran Lepidopleurus ker­ Summary of zonation on transects guelensis and Hemiarthrum setulosum occur under The basic pattern of zonation at Macaroni Bay boulders, the former being more common. Pelecy­ Transect I is common to all stable plane sur­ pods Kidderia sp. were not noted here, in contrast to faces that are not shaded and are exposed to the apparently similar conditions at Transvaal Cove. open sea. However, much of Marion's littoral Actiniarians are common against the substrate and environment is made up of surfaces that are the asteroid Anas1erias rupico/a is fairly common. frequently shifting or subject to abrasion (boulders of The pycnogonid Tanys1ylum cavidorsum is rare, as is Transvaal Cove) and are shaded. face the substrate, the sipunculid Go/fingia ohlini. Nemerteans are fairly or are protected from the influence of the open sea common. A variety of polychaets inhabit the detritus, (Dlliker's Point and Ship's Cove). It is on such the most conspicuous being Platynereis australis. surfaces that major departures from the basic pattern Other common species include Syllis variegata, may be noted. Table 3 summarizes the relevant Phyllodoce pO~I'phyl/a, Cirratulus cirratus and differences.

Plate 6. Duiker's Point Transect locality. Fronds of Durvillea antarctica diminish the force of the swell. Kerguelenella lateralis

Lumbricil/us spp. Tide pools occurs in small numbers. Hemiar1hrum setulosum Tide pools of any extent are uncommon on MariOn's may also be found under boulders but is far more youthful shores, those that do occur being generally abundant on Phymatolithon sp. Nacella delesserti is Verrucaria sp. small and shallow. Salinity and temperature varia­ surprisingly rare in this pool, though there are many ~- I tions increase with increasing distance from the sea juveniles on the stipes of Desmarestia rossii. A Rhodoglossum sp. and the distribution of organisms reflects these pelecypod Philobrya sp. and the minute gastropods I._._..., variations. Salinity and temperature of three pools at Laevilitorina caliginosa and Eatoniella kerguelenen­ Rhodymenia spp. varying distances from the sea at Paddy Rocks sis also occur on algae. E. kerguelenensis Jives under (Transvaal Cove) have been referred to (Table 2). A stones as well. Echinoderms are poorly represented, description of these pools and their biota follows: only the tiny ophiuroid A mphiura tomentosa Enteromorpha1-.J bulbosa Poo/1: This large, roughly circular pool, about 4 m (synonym Nullamphiura marionis) and the asteroid Porifera in diameter and 1-2 m deep, has direct communica­ Anasterias rupicola being recorded. tion with the sea by a lava tunnel about 8 m long, 3 m Ascidians are rare and cryptic, inhabiting the sides wide and 2 m high. The pool enjoys calm conditions and under-surfaces of boulders. Two species Core/la for most of the time, but is swept by large swells at eumyo1a and Polyzoa reticulata were collected. least once a month when swells of 2 m or more invade Sharing this niche are the sponges Sycon kerguelen­ Paddy Rocks. Breakers of more than 4 m scour the sis. Leuconia joubini. and Ectyomyxilla chilensis. whole area a few times during the year. In spite of P lanarians are common on algae while a few this, the general appearance of biota in the pool is one bryozoans and hydroids were collected on the sides of of luxuriance due to shelter. Investigations were boulders. Nemerteans, sipunculids and holothurians cm carried out by snorkel diving. The floor of the pool is occur under the boulders. 136 strewn with boulders and smaller rocks encrusted A rich polychaet fauna inhabits crevices, the under­ together by lamellate disc-forming Phymatolithon sp. surfaces of boulders and substrate detritus. The as well as species of Schizoseris and Heterosiphonia. following species were recorded: Harmothoe berg­ Other common algae include the large branching srroemi, Tharyx sp .. Phyllodoce po/yphylla, Platy­ Desmarestia rossii and [). chorda/is. Encrustations of nereis austra/is, Haploscolopos sp., Polycirrus ker­ Codium sp. occur from the upper sub-littoral to guelensis, Potamil/a antarctica and Thelepus setosus. deeper levels. Aggregations of Adenocystis utricularis The most conspicuous of these is Platynereis australis carpet the floor in parts where Ballia callitricha is not whose matted tubes occur on the under-surfaces of uncommon. Schizoseris ?kerguelensis and Lophurel­ boulders as well as in detritus. Serpulids (Spiror­ /a sp. grow fairly commonly from the sides of binae) are also quite common at LWS particularly 14------770cm - ~ boulders in the upper sub-littoral. Helicosiphon sp. and Paralaeospira levinseni. L Bostrychia spp. • Rhodymenia, ~ Lithothamnia Various amphipods, mainly juvenile, inhabit the Pool 2: This small pool, not more than a metre in • Common Cladophora turf ~ Ballia callitricha algae. Many specimens of Cerapus ?tubularis were diameter, is less than 30 cm deep and is replenished ~ Rama antarctica • Fairly common also collected. The isopod Dynamenella huuoni is by splash with every tidal cycle. It is flopred by dense ~lea antarctica - Rare fairly common and the tanaid rare. .. Verrucaria spp. Anatanais gracilis encrustations of Lithophyllum sp. which never ~ A single brachiopod was found under a stone where extend above the water level. Planarians crawl over the tiny amphineuran Lepidop/eurus ke~guelen sis the Lithophyl/um in numbers and the isopod Fig. 7. Duiker's Point Transect. The distribution a nd relative abundance of the major tone-forming organisms. 22 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J . Antarct. Res., 1976, Suppl. 1 23

Table 3 The greatly reduced species diversity of Pools 2, 3 and plants under slightly increased exposure, and the Cape Davis pool reflect the specialized conditions may span 10 m in vertical extent to form a Variations in species distribution at the various transects found there. The disappearance of lithothamnia, and continuous curtain of holdfasts, stipes and the presence of some or most of the following species fronds under extreme exposure (Cape Davis). PHYSICAL MACARONI MACARONI SHIP' COVE TRANSVAAL DUIKER'S indicate widely fluctuating temperature and salinity: 8. Lithothamnia reach their upper limit a few FACTORS BAY I BAY 2 COVE POINT Enteromorpha bulbosa, Rama anrarctica, Bosrrychia centimetres below the upper limit of Durvi//ea Substrate Unbroken, unbroken. Boulders. Mean diam. 215cm Boulders; mean Boulders. mean spp., Rhizoclonium riparium, Tigriopus angularus holdfasts under calm conditions, a metre or so fairly smooth smooth diam. 68cm diam. 56cm and Lasaea consanguinea. below Durvillea under increased exposure (Cape Hooker), and several metres below under Approximate ave- The pools of Cape Hooker extreme exposure (Cape Davis). rage slope in About 1,5 km west of Cape Hooker 20-30 m high 45° !5° !5° 150 so 9.Po()•siphonia, Schizoseris, Lophure//a and My­ degrees to the cliffs occur some I 00 m from the sea. Along the cliff riogramme frequently form a distinct belt (or horizontal bases very large boulders (more than a metre in belts) in the upper half of the upper sub-littoral Relative expo- diameter) extend for about 20 m. These are followed under calm conditions (data lacking for exposed sure to wave Moderate Slight Moderate Slight Moderate by an area of low, flat reefs and rock systems which conditions). action interconnect over a breadth of about 60 m to form a 10. Macroscopic invertebrates are cryptic above the system of deep channels, gullies and pools. The upper limit of Durvillea except under conditions system is protected from much of the sea's force to of extreme exposure (Cape Davis) where ZONATION Exposed Sheltered or landward but becomes progressively exposed to Kerguelenella lateralis may form dense aggrega­ surfaces occluded seaward. Detailed investigation of this area resulted tion in the region of HWS and Lasaea surfaces in the conclusion that the characteristic heavy seas of consanguinea may attach itself to exposed the south coast enable organisms of the lower and Verrucaria Verrucario Verrucoria Verrucaria Verrucoria surfaces on top of cliffs. Lichen zone Verrucaria sub-littoral zones to extend into pools, some even to a Ulorhrix Ulorhrix Kerguelenella Kerguelenel/a Ulorhrix Bosrrychia Kerguelenella Kerguelenella Ulorhrix Kerguelenella Rama considerable distance from the sea, e.g. lithothamnia, Sub-littoral of Gunner's Point Porphyra Kerguelenel/a Codium sp., Durvillea antarctica, Leptosomia sp., Distribution of organisms to a depth of 4 m on the Hemiarthrum setulosum, Nacella delesserti and south-south-west sub-littoral cliff face of Gunner's Porphyra zone { 'fmhrix Verrucaria Porphyra Verrucaria Ulorhrix Vernlt"aria serpulids. Point (on the northern border of Transvaal Cove) Porphyra Ulorhrix Ulorhrix Bosrrychia Porphyra Rhodymenia sp.2 Simpson79 ( 1976) records that lack of Durvillea was investigated. Kerguelenella Porphyra Kerguelenel/a Rhodymenia sp. I Cladophora cover is limiting to H. setulosum. It was therefore Below the characteristic algae of the upper sub­ Rhodymenia sp. I Kerguelenella Rama interesting to record its abundance on dense littoral, a continuous carpet of Coral/ina sp. extends Kerguelenefla Kerguelenella lithothamnia in Durvillea-free pools of this area. for 1-2 m in the sub-littoral to the virtual exclusion of other forms, a few Desmarestia rossii and D. Kelp zone Dun•illea Dun•illea Durvillea Bosrrychia Urhorhamnia Durvillea Zonation on steep stable plane surfaces chorda/is only being noted. Species diversity increases Lithothamnia Lithothamnia Lirhothamnia Abroreiaf LRptosumia Lirlwthamnia Much of Marion's rocky shore consists of vertical or with increasing depth and the cover of organisms Hemiarrhrum Leprosomia Hemiarrhrum Verrucaria Nacella Hemiarrhrum very steep cliffs on which it is extremely difficult to against the substrate is complete. Typical algae down Nacella Kerguelenella Nacella Nacel/a Hemiarrhrum work. Observations at points of varying exposure to 2 m include Lophurella hookeriana and species of Nacella around the island allowed the following general Desmarestia, My riogramme, Schizoseris and Hetero­ conclusions to be drawn: siphonia. while below this (to 4 m) Polysiphonia Lower Red zone Lirhorhamnia Urhorhamnia Lirhorhamnia Lirhorhamnia Rhodoglossum Bollia 1. Porphyra increases in abundance with increased urbanoides and Geminocarpus geminarus occur as Schizoseris Schizoseris Durvil/ea Po~rsiphonia Lirhorhamnia Hemiarrhrum exposure (Cape Hooker) up to a point beyond Desmaresria Myriogramme Po~rsiphonia Schizoseris Po~l'siphonia Nacella well. (Data Desmaresria Schizoseris Myriogramme Schizoseris which extreme exposure (Cape Davis) limits Hydroids are abundant against the cliff face at incomplete) Myriogramme Myriogramme growth. Vertical faces are less favourable than depths of 1-2 m, particularly Symplectoscyphus Nacel!a slopes. marionensis and Sertularella picta. Dense aggrega­ - 2. Bostrychia spp. favour south facing slopes or tions of tubularids occur under shelves and Exosphaeroma gigas is quite common. Amphipods measured, the greater diversity of organisms suggests conditions of shade and freshwater seepage. overhangs. An interesting find was the stauromedusan Pontogenia sp. inhabit the substrate and also swim in greater stability of this parameter here. 3. Rama antarctica frequently grows in association Haliclystus anrarctica, previously known from South mid-water. Abundant growths of the filamentous Rhizoclo­ with Bostrychia (north of Transvaal Cove) but Georgia and Antarctica. At Gunner's Point it was Pool 3: This is further back from the sea than Pool nium riparium occur around the edge about 20 cm may also occur in abundance high up on wave­ taken from Po(ysiphonia urbanoides at 4 m depth. 2 and similar in size. As a result of infrequent above water level. This alga forms a belt 5 cm wide lashed cliff faces. (Goodhope Bay, Cape Davis). The bryozoan provisionally identified as Menipea splashing it suffers wide fluctuations of salinity and and is overtaken towards its lower limit by Bostrychia 4. Rhodymenia species 1 does not grow on shores crispa is fairly common under sheltered conditions at temperature. Lithothamnia are entirely absent, their mixta which extends below water level. Rama of greater inclination than about 45°. 3 m, where the asteroid Anasterias rupicola occurs as place being taken by luxuriant growths of Entero­ antarcrica grows around the sides of rocks in the pool 5. Rhodymenia species 2 and the associated Clado­ well. At this depth a wide variety of organisms occurs morpha bulbosa and Rama antarctica. The Entero­ and does not extend above water level. The tangled phora and Rama which together form a turf among the algal fronds, particularly the amphipod morpha is submerged, but Rama grows on the sides filaments of this alga are inhabited by large numbers complex favour a rough substrate (north of Jassa falcata and the isopods Dynamenella hurtoni of the pool both out of the water and in it. The tanaid of Anatanais gracilis, amphipods and copepods. Transvaal Cove) where its colour is predominant­ and Jaeropsis curvicornis. The tanaid A natanais Anatanais gracilis and the pelecypod Lasaea consan­ Anatanais extends above water level in the filaments ly dark brown. It also grows in abundance as gracilis is fairly common. Also noted were nemerteans, guinea are common among the filaments of Rama. of Bostrychia and Rhizoclonium. The isopod pale green sheets covering many metres in the pycnogonid Tanysty/um cavidorsum, holothuroi­ The harpacticoid copepod Tigriopus angu/atus lives Exosphaeroma gigas is common, and planarians vertical extent on steep smooth faces of extreme deans and a single specimen of the hirudinean among the fronds of Enteromorpha in abundance. ?Procerodes sp. crawl in numbers over the sandy exposure (Goodhope Bay, Rook's Bay). It is Antarcticobdella crozetensis. The pelecypod Lissarca Planarians and amphipods are rare and isopods bottom and on the under-surfaces of stones. absent from intermediate conditions. rubrofusca is fairly common on the alga Polysiphonia altogether absent. Oligochaets Lumbricillus sp. and insect pupae occur 6. Abroteia/ Verrucaria becomes abundant under urbanoides. Several features of Pool3 were also noted in a pool against the substrate and among the filaments of conditions of extreme calm (Transvaal Cove) 10-15 cm deep on top of the coastal cliffs of Cape Bosrrychia and Rhizoclonium. where there is no competition from Durvillea Boulder shores Davis. The fact that the pool was situated some :30 m It may be concluded that the biota of Pool 1 are antarctica. Trypot Beach: The beach at Trypot is composed of back from the cliff edge is indicative of the intense typical of the shallow sub-littoral rather than of the 7. Durvil/ea antarctica is absent from calm faces small boulders (6-10 cm diameter) and is inhabited wave action at this point. Though salinity was not widely fluctuating conditions higher on the shore. (Transvaal Cove), occurs as a single row of for parts of the year by King penguins and elephant 24 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 25 seals. Even during calm seas the boulders are shifted and under holdfasts of kelp has been mentioned in and rolled by each swell so that their overall earlier se.::tions. After storms much seaweed could be Firsr £rophic level appearance is smooth and devoid of life. However, a found in piles on the shore, the most conspicuous Verruc:aria Brown algae Ulothrix POfphyra Durvillea Diatom film Phytop4ankton (Primatv producers! surprisingly rich fauna consisting mainly of the light . being Macrocystis pyrifera and Durvillea antarctica. brown amphipod Hyale grandicornis occurs among This provided an opportunity of investigating more the boulders between tide marks. Hyale inhabits pock fully the fauna associated with large algae. -~----L-\::· ~------i----;,,,.;;~-; marks in the boulders from about the undersurface of Durvillea antarctica: The fronds and stipes of ~ ~ ~ Polychaets ~ t ~ copeooml ~ / ~ ~Sp o~l ./ \ Ga1mardia K·d•- . the second superficial boulder to deeper levels. Hya/e (011barod (Kerguelenella) Ploty· A h' d Nacena / - ' ~mna lAmphipoda: Hyale hirtipalma has formed irregular boulders of widely varying sizes Bryozoa: Menipea crispa, Tubulipora sp. (provision- which lie piled chaotically along the base of the cliffs. al identifications) Fig. 8. Littoral food web of Marion Island. At the time of observation very large swells Pelecypods: Philobrya sp., Kidderia oblonga approached the shore from the south-east. Distribu­ Gastropoda: Nacella delesserti tion patterns of organisms suggested that swells of pa11 of its diet. primarily of Rhodymenia species 2. Holothuroidea: Pseudocnus laevi~atus this nature were normal for this part of the coast. Macrocystis pyrifera: Along most of the east coast The Sheathbill Chionis feeds on Hyale spp. from Kerguelenella lateralis is present in numbers, and Of great interest was the observation of Durvillea extensive beds of Macrocystis occur some lOO m among fronds of Porphyra and frequently ingests the orange mite (Mesostigmata) abundant. Just antarctica growing on large boulders higher up on the offshore. On the exposed south and west coasts pieces of Porphyra as well. above the Durvillea antarctica zone rich growths of shore than luxuriant carpets of Porphyra and Macrocystis is far less common and occurs further Notothenia corriiceps has a wide variety of food Rhodymenia species I occur and together with Rhodymenia. The popular practice of using organisms Porphy ra form a distinct belt. Below the Durvillea offshore and in patches. organisms. At times their stomachs were found as indicators of effective tidal heights causes grave lithothamnia encrust the rocks and Nacella delesserti The most conspicuous organism associated with distended by great quantities of calanoid copepods, problems in such cases since the lower littoral would may be found quite commonly on it. Macrocystis is the pelecypod Gaimardia trapesina chaetognaths and hyperiid amphipods. Anasterias have to be assigned a higher level on the shore than the Tiny pools form in depressions on the boulders, which attaches itself to the floating fronds in rupicola was frequently found feeding on Nacel/a upper littoral. The importance of substrate stability for and the contention that there is persistent strong considerable numbers. A count of 71 speciments was delesserti. The gull Larus dominicanus takes a very large slow-growing algae cannot be emphasized too wave action on this coast was born out by the obtained from a random collection of 22 fronds. A heavy toll of Gaimardia trapesina which grows much, and these anomalous situations are discussed in attached to the fronds of offshore Macrocystis. presence of abundant copepods and Enteromorpha in a later chapter. bryozoan encrustation on Gaimardia shells was provisionally identified as Hippothoa hyalina. The Simpsonso ( 1976) has published interesting data on such pools more than 10 m vertically above the upper The boulders here vary from about 30 cm in serpulid polychaet Romanchella perrieri is common food preferences of littoral organisms on Macquarie limit of Porphy ra. diameter to more than one metre. Frequent and on all mature Gaimardia as well. Island, but most of these await verification under A beach situated about lOO m east of the above irregular shifting results in further disruption of the The fronds of Macrocystis are further colonized conditions prevailing at Marion and are not included location was examined. Composed of boulders of a zonation pattern. Boulders recently rotated or cast up by abundant hydroids, especially Obelia geniculata in Fig. 8. size comparable to those of Transvaal Cove on from a deeper level are colonized by organisms whose and Silicularia rosea. A list of fauna associated with Marion (about 50-70 cm diameter), the beach is growing season occurs at that time. It would seem that the much branched holdfasts follows: The m toral ecology of Prince Edward Island subjected to very heavy surf. Durvillea antarctica is Porphyra sp. grows vigorously in early and mid Hydrozoa: Myriothela meridiana, Sertularella picta During April 1973 a one-week visit was paid to the entirely absent. Boulders bearing bleached white summer and that Rhodymenia species I colonizes Nemertea: Tetrastemma sp. (provisional identifica- little-known Prince Edward Island. Lying just 22 km lithothamnia and serpulid tubes lie scattered through­ new available surfaces primarily in late summer and tion) north-north-east of Marion, Prince Edward is rarely out the tidal range. An interesting feature was the autumn. Large rough-textured boulders high up on Polychaeta: Thelepus ?setosus landed on and has no buildings or amenities of any absence of Porphyra, its level on the shore being this shore are well covered by Rhodymenia turf and a sort. taken over by a luxuriant belt of Rhodymenia species I. short green alga. Kergue/enel/a lateralis juveniles and Bryozoa: Crisia sp., Tubulipora sp. Ophiuroidea: Amphiura tomentosa, Ophiacantha Observations and collections were carried out in adults are common. Nearer the sea scattered vivipara, Ophiurolepis martensi the intertidal zone to the east of a 500 m high Porphyra may be found together with luxuriant escarpment which divides the island. Prince Edward growths of Rhodymenia species I and a little Food web is roughly rectangular in shape and its long axis is Enteromorpha bulbosa. Boulders of the lower littoral orientated along a north-west direction (Fig. 9). true! bear encrustations of lithothamnia and other red During the course of the investigation of the littoral north I, many observations on feeding behaviour could be \Illest algae. Durvillea is present on large boulders and North-east coast: A few hundred metres west of Fbint; smaller ones that are firmly wedged between other made, and a food web diagram (Fig. 8) was compiled. However, certain links should be regarded with where Albatros river flows into the sea very large boulders. Nacella delesserti was noted on the boulders (l-2 m diameter) lie strewn along the shore. lithothamnia, but it was not possible to approach caution since they are based on only one or two observations and may not therefore represent a On the landward faces of these boulders several close enough to check for Hemiarthrum setulosum. metres back from the sea Rama antarctica and ~ntA Nacella extends up the shore to higher levels than on typical pattern. In this respect mention should be made of Durvillea being grazed by Nacella. In a few Bostrychia sp. are quite common. Oligochaets and calmer coasts and could be found at levels reached by insect pupae inhabit the filaments of Rama. Below 1 2 3 Rhodymenia turf. instances heavy radula strokes on broad Durvillea ki~t!.is I ' fronds were noted. These could only have been made these growths patches of Verrucaria encrust the rock, A brief investigation between and underneath and share the same level reached by Kerguelenel/a boulders revealed abundant serpulids Romanchella by Nacella but how frequently Nacella indulges in Durvillea is not known. lateralis. Mites (Halozetes sp.) are abundant on the sp., amphipods, and a few Nacella de/esserti and Verrucaria where it attains continuous cover. On the South Cape Kerguelenella lateralis. In other cases the same food organisms for certain species were noted time and again. Stomach contents boulders closer to the sea Porphyra becomes quite Kelp fauna of the plunder fish Harpagifer bispinis showed that common and is succeeded at lower levels by small Fig. 9. Outline of Prince Edward Island showing localities The occurrence of organisms on the fronds, stipes isopods, amphipods and polychaets form a major quantities of the turf-like complex composed mentioned in the text. 26 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 27

East coast: Observations were carried out immedi­ South coast ~ ~ ...... , ~~ ..:...·- ately south of R.S.A. Point where very calm Exposure increases along the south coast moving conditions prevail. Here stratified tuff at the base of a westwards and this is reflected by the increasing scoria cone forms a soft substrate for colonization by vertical range of Durvil/ea antarcrica and the intertidal assemblages. This section of coast has appearance of extensive Rhodymenia species 2 turf numerous deep crevices and gullies which dissect the carpets on vertical faces in the region of South shore (nearly at right angles to the coastline) to a Cape. Shortage of time and rough seas precluded depth of 10 m or more. Many are very narrow (2~50 cm) detailed sampling. and taper to their ends. Swells approach the entrances to the gullies obliquely and travel along Conclusions them until their energy is confined at the landward As far as could be ascertained in the time available, end where they splash vigorously and send spray the littoral ecology of Prince Edward follows very cascading onto the level substrate on either side. The closely the pattern noted on Marion. collective splashing of the numerous gulleys results in an extensive area of frequent inundation. This area is colonized by a continuous carpet of luxuriant turf comprising mainly Rhodymenia species 2 and DISCUSSION Bostrychia mixta. Generally golden to light yellow in colour it darkens when occurring in a cleft or depression. Also common are patches of Enrero­ Factors influencing distribution of morpha sp. No lichen zone is apparent. orgamsms A great upward extension of Durvillea antarctica Tides and lithothamnia may be noted in the crevices at their The tidal regime on Marion has been described above ends where the splash is more vigorous. At such (p. 16). The tides are of the regular semi-diurnal type points Durvil/ea may extend to near the edge of a and have a range of 71 cm at spring tides. This small crevice and three or four holdfasts may occur amplitude is the lowest recorded in the sub-Antarctic, Plate 7. Configuration of gully and wave action result in splash in mechanism which raises effective tidal height by 5 m vertically above one another. To seaward along the being 34 cm less than the spring range at Tristan da immediately below observer. walls of the crevices the upper limit of Durvil/ea is Cunha (Baardseth,8l 1942). displaced downwards together with lithothamnia and From early times workers have recognized that organisms at Macquarie Island. Taking measure­ of violent storms and prolonged periods of calm have the holdfasts occur singly. A narrow belt of zonation on the shore is not caused solely by tidal rise ments from their figure and using upper limits of been presented earlier (p. 6). Rhodoglossum may be distinguished just above and fall but by a host of additional factors which dense growth of the organisms, Durvillea reaches As would be expected the most profound effects of Durvillea's upper limit where it grows among turf influence the degree and duration of wetness at a near MSL at one of their transects and above MSL at severe wave action may be noted on the exposed west carpets of Rhodymenia. The turf carpets themselves particular level on the shore. These factors may vary another. and south coasts where the size of the waves exceeds extend over the edge of the crevices and grow down widely over short distances and include such variables Arnaud92 (1974)' has raised the interesting point the tidal range by several metres. The large waves, the walls in abundance particularly to seaward where as slope of shore, nature of substrate, intensity of that the fronds of Durvillea antarctica at Kerguelen coupled with frequent onshore gales, have the effect Durvillea's upper limit is displaced downwards. wave action, degree of exposure and climate. The Island hang down and float on the surface of the of extending the marine influence well inland. Yellow, Lithothamnia in parts extend above the upper limit formidable task of expressing the effective wetted water when the holdfasts are above sea level and that white and grey lichens characteristic of the upper of Durvillea well concealed by Rhodymenia turf. area with every tidal cycle in physical terms has been the fronds float up to the surface when the holdfasts supra-littoral on the east coast (e.g. Transvaal Cove) Deep within the crevices below the lower limit of pointed out by Lewis82 (1972) and littoral ecologists are below sea level. Where the whole plant is above extend more than 150 m above sea-level at Triegaardt Durvillea a lower red algal zone is absent, the rocky have preferred to use indicator organisms to express sea level it is at least drenched by frequent splash at Bay and may occur several hundred metres inland. surface being covered only by abundant lithothamnia. the different effective tidal heights (e.g. Doty,83 1946; all states of the tide. He concluded that Durvillea Small fronds of Porphyra occur on particles of The Rhodymenia turf supports huge populations Stephenson & Stephenson,84 1949; Womersley & should be referred therefore to the "infralittoral" volcanic ash 50 m inland on top of 20-m high cliffs of the pelecypods Kidderia bicolor and Lasaea Edmunds, 8s 1952; Lewis,86 1972). This policy has also (sub-littoral). On Marion this is the case in many but between Cape Davis and Mixed Pickle Cove. consanguinea as well as oligochaets Lumbricillus been employed in the sub-Antarctic and Knox81, 88 not all instances, particularly at Macaroni Bay Only the hardiest organisms can survive on the macquariensis and L. aestuum. Also common are (1960, 1968) has drawn up useful summaries of Transect 2 where both fronds and holdfasts were seaward faces of exposed cliffs. The lush carpets of amphipods Hyale sp. and the tanaid Anatanais zonation patterns using indicators on sub-Antarctic above sea level and received no splash for extended Rhodymenia turf and the occurrence of Durvillea at gracilis. shores. His terminology has been followed by periods during low tide and calm seas. Goodhope Bay provide good examples. A similar The amphineuran Hemiarthrum setulosum is fairly Chamberlain89 (1965), Chamberlain, Holdgate & The difficulty of using indicator organisms is zonation pattern was noted at Mixed Pickle Cove common on lithothamnia where a few Nace/Ja Wace9o (in preparation) and the present writer. further illustrated by the disappearance of Porphyra where a continuous carpet of algal turf spanned more delesserti and Kerguelene/Ja lateralis could be noted It is instructive to note the level reached by on Marion in the middle of the winter. than 5 m above the Durvillea. as well. Durvillea antarctica on three of the transect shores at In conclusion it is the writer's opinion that levels Cliffs that are not vertical allow some of the swell's Boulder beach: Further observations and collect­ Marion. Durvillea is normally regarded as occurring reached by organisms on sub-Antarctic shores should force to be dissipated, and scant growths of Porphyra tions of organisms were carried out at a boulder­ in the lower littoral reaching to L WN. If the upper be expressed as centimetres above a small perennial of and Rhodoglossum can survive above extensive strewn beach a few hundred metres south of R.S.A. limit of dense Porphyra growth is taken as widespread sub-Antarctic occurrence that occupies curtains of Durvillea (Cape Davis). An almost Point. All the typical zone-forming organisms from delineating HWS and the upper limit of delicate the lowest level that can be conveniently reached. continuous deluge of spray permits the tiny the lichen zone to the lower red zone could be noted. mixed red algae below Durvillea as marking L WS, Algae such as Polysiphonia or Myriogramme would pelecypod Lasaea consanguinea to colonize rocky Between the boulders and against the substrate a rich then Durvillea reaches to just above effective L WN at probably fit the requirements suitably. Once sufficient faces some metres back from the cliff edges at such and varied fauna was encountered. A species list Macaroni Bay Transect I, to well above MSL at data throughout the sub-Antarctic had been gained points. follows but is not complete. Macaroni Bay Transect 2, and to just below MSL at one would be in a better position to define effective Boulders of comparable diameter to those of the Pelecypoda: Lasaea consanguinea Transvaal Cove. Clearly wave action and degree of tidal heights more precisely. Transvaal Cove Transect (50-70 cm) are devoid of life Gastropoda: Nace/la delesserti, Laevilitorina caligi- slope are playing a major role here, and to retain Plate 7 illustrates an extreme example of altered as a result of continuous shifting <\nd rotation nosa, Kerguelenella lateralis Durvillea as an indicator of L WN, Porphyra would effective tidal height due to the combined influence of (northern border of Triegaardt Bay). Platyhelminthes: ?Procerodes sp. have to be abandoned as a major indicator of HWS topography and wave action. On the east coast the effective tidal range at each of Polychaeta: Romanche/la sp., Platynereis australis in favour of an organism growing at higher levels. the transects examined far exceeds the astronomical Isopoda: Exosphaeroma gigas, Munna sp. Kenny & Haysom9l (1962) have presented data (p. Wave action tidal range (Figs. 3-7). During the rare occurrence of Amphipoda: Hyale grandicornis, Pontogenia sp. 258, Fig. 9) showing levels reached by indicator Data on the average size of swells and the periodicity strong onshore winds large quantities of algae are '1

28 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 29

Plate 8. Calm conditions on ca~t coa~t. Floating fro nd~ of Macroc:rMi' clearly visible off,hore. The furthc>t point is Plate 9. Rough condition' on cast coa>.l. Prince Edward bland lies 22 km to the north-east. East Cape. torn loose from the substrate by the resultant swell zonation patterns also. A north-east slope off mechanism may be set up. A spectacular example of occupies its normal level on east coast shores with and wash up onto boulder beaches. The most Gunner's Point (Transvaal Cove) had well developed this occurs just south of Duiker's Point (Plate 7). fronds floating on the surface of the water and vulnerable are Macrocystis pyrifera, Durvillea ant­ Porphyra and Durvillea zones while 3 m away on an Waves are confined in a small gully where at the holdfasts being momentarily exposed between swells. arctica and delicate rhodophytes. Boulders from east-south-east slope Durvillea was entirely absent landward end even small ones slap the rock with Mention has been made in an earlier section of the deeper levels may be cast up in large numbers so that and Porphyra much reduced. On a south-south-west considerable violence. A jet of coarse splash drenches deep gullies dissecting the stratified tuff shore on the beaches become checkered with bleached lithotham­ slope at the same point, Porphyra was alJ but absent an 8 m rocky face with great regularity to raise the east coast of Prince Edward Island. Here an extensive nia-covered rock. Conversely, no adverse effects due and there was no Durvillea. Swell direction in this effective tidal height at once by about 5 m. Durvillea area is subjected to frequent splash as the swells to periods of prolonged calm were noted. The high locality is generally from the north-east and the faces anrarctica reaches its upper limit very near the top of "erupt" from the confined landward ends of the humidity, relatively constant low temperature and with southerly aspect enjoy conditions of relative the 8 m rocky face and the fronds hang free with only gullies. This frequent inundation and the gradual almost continuous cloud cover preclude mortality calm. their tips reaching the surface of the water. Ten slope of the rock between the gu llies results in a due to desiccation. Inclination: The lesser the degree of inclination, the metres away on either side of this point where the continuous carpet of luxuriant Rhodymenia turf Plates 8 and 9 illustrate two extremes of slower the drainage of the surface. In this connection splash mechanism is not operative, Durvillea (species 2). conditions encountered on the east coast. it is interesting to note the luxuriant development of Durvillea and Leptosomia ?simplex at Macaroni Bay Topography (aspect, inclination and configuration) Transect 2. Not 30 m away at Macaroni Bay Transect Aspect: Shores facing every point of the compass are I, Leptosomia is entirely absent and Durvillea less encountered on an oceanic island such as Marion and well developed. changes of aspect bring very few environmental Furthermore, the more gradual the inclination, the variables into play. This may be contrasted with greater is the surface area between effective tidal conditions on continents where changes of aspect heights for colonization by intertidal organisms. may involve great distances incorporating changes of While unbroken shores of gradual inclination are rare climate, hydrology and bedrock (Lewis. 93 1972). The on Marion, boulder-strewn shores of this nature are oceanic climate enjoyed by Marion with its high common. Drainage from substrate is further impeded degree of cloudiness may further result in there being by the presence of the boulders and smaller particles. little significant difference between insolation on the This, together with general protection provided by north a nd south coasts. Until figures on localized the boulders, allows an accumulation of mixed climate over long periods on Marion's west coast detritus against the substrate to form a relatively become available it is not possible to predict the stable niche harbouring many forms of invertebrates effects of differences in rainfall, humidity and similar (for example Duiker's Point Transect). variables. It is certain however that one of the most Vertical shores drain almost instantaneously, and important single factors is the high incidence of are very lean in biota above the upper limit of onshore gales on the west coast. These winds may Durvillea on the east coast. The almost continuous blow over a fetch of 8 000 km and generate swells of deluge of spray on the west coast however results in colossal magnitude. Only organisms adapted to with­ the lush carpets of Rhocz\'menia above Durvillea on stand a battering of the severest nature can survive on vertical shores at places such as Goodhope Bay. these west-facing shores. Plate I0 illustrates typical Configuration: Configuration of rock at a particu­ conditions on a south-west facing shore. lar place may result in a wave's energy being Very localized changes of aspect result in different funnelled into a confined space where a splash Plate 10. ·1 he exposed south coast showing breadth of surf zone. View towards the south-east from Good hope Bay. 30 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 31

Substrate be colonized primarily by Rhodymenia turf in the The nature of the substrate is an index. of Porphyra zone with very little Porphyra. It may be topographical micro-variations and will accordingly that shady conditions influence this occurrence. influence the speed of drying out of a rocky face. On A stratified tuff shore on Prince Edward's east the unbroken shores of Macaroni Bay crevices and coast was the softest rock examined and could readily grooves in the rock retain water and are frequently be scored with a knife. The upper littoral here was colonized by algae such as Rama antarctica and colonized largely by algal turf. Bostrychia sp. Living among the fronds of these algae On broken shores the nature of the substrate are the pelecypod l..Asaea consanguinea and the changes over a very small vertical range. Natural oligochaets Lumbricillus aestuum and L. macqua­ sorting of boulders during heavy seas results in a riensis. Higher on the shore striations are favoured by gradation of particle size from large in the superficial the mites Halozetes marionensis and the orange mite layers to small against the immovable substrate (Mesostigmata). According to Simpson94 (1976) beneath. A stable boulder shore breaks the force of Kerguelenel/a lateralis is weakly attached and swells of average size and a great variety of inhabits crevices for protection and to reduce invertebrates inhabits the deeper levels between desiccation. On Marion's sheltered shores this species boulders where they find protection under stones, on is confined to depressions. On exposed shores, the sides of stones and among detritus. Common however, it may be found high up on open surfaces organisms of these niches include actiniarians, well above the level where waves could dislodge it, platyhelminths, nemerteans, sipunculids, polychaets, but where an almost continuous deluge of spray isopods, amphipods and a wide variety of molluscs ensures constant hydration. (Duiker's Point and Transvaal Cove Transect). Lower down on unbroken shores narrow crevices Shores composed of very small boulders (Trypot are colonized by large numbers of pelecypods Beach) are understandably lean in species, and those Plate 12. Um.table ' ub, tr:uc. Frc4ucnt rotat ion of boulder. results in une \ cn dJ ,tribution of organi' m'. :>. otc Kidderia bicolor and K. oblonga. that do occur tend to congregate at the deepest levels. absence of Dunill<'a amarct1ca (l ranwaal Cm e). At certain points (Ship's Cove and near Macaroni A notable exception is the amphipod Hyale Bay) lava of the more recent eruption has an grandicornis which inhabits pock marks in the stones extremely rough texture. Such surfaces appeared to and probably avoids being crushed in this way. The importance of substrate stability becomes meteorological station). Pools of reduced salinity shore (south of Soft Plume River). This has a apparent on shores composed of boulders of greatly harbour rich growths of Enreromorpha bulbosa. and stabilizing effect on broken shores thus allowing varying size. South of Kildalkey Bay Durvillea frequently there are abundant copepods Tigriopus optimum development of sub-fauna. However during antarctica was noted growing on large boulders at angulatus. heavy seas the tough fronds of Durvillea whip back higher levels on the shore than Porphyra and The high degree of cloudiness and high humidity and forth over rocky faces causing abrasion, and not Rhodymenia which had colonized smaller, less stable reduce evaporation and it is likely that this allows many organisms occur in the open. Notable boulders. Such occurrences add to the difficulties of organisms that are prone to desiccation to extend exceptions are the tiny amphincurans 1/emiarthrum defining effective tidal heights by means of indicator higher up the shore. setulosum and juvenile Nacella delesserti. Encrusting organisms. The strong winds have been referred to and the far lithothamnia can survive in abundance beneath dense Plates 11 and 12 illustrate a stable and unstable reaching effects of this single factor cannot be over­ curtains of Durvillea but delicate algae such Ballia boulder shore respectively. emphasized. Schultze95 (1971) analysed five years of callitricha, Schizoseris spp. and Polysiphonia are The sandy beach at Ship's Cove is devoid of wind direction data and found that the winds have a confined to LWS at which level the Durvillea fronds macroscopic forms of life and is periodically eroded westerly component 78% of the time and that are always borne up by water. It is interesting to note away to expose underlying boulders. conditions of calm prevail only 8% of the time. Gales in this connection that the upper limit of lower red There are very few places on Marion where occur on average on 107 days per year. On the west algae coincides with the upper limit of lithothamnia proximity of sand or grit affects zonation. One of coast salt water in the form of spray and splash is on boulder 3 of the Ship's Cove Transect where these is the eastern shore of Good hope Bay where grit borne inland over the cliff edges to such an extent Durvillea is absent. scours the exposed faces of embedded rock and that Porphyra may be found alongside maritime Colonization of algae: An important feature of removes all forms of life. halophytes such as Tillaea moschata. littoral ecology on Marion is the rich and varied Conversely, on the east coast the strong winds fauna that lives in close association with lichens and Climate transport spray from broken waves back to sea so algae. In the upper supra-littoral various dipterans A description of climatic conditions has been given in that the spray zone is greatly reduced. However during and collembolans inhabit piles of rotting kelp and an earlier section (p. 7). Marion and Prince Edward are heavy seas when waves reach higher levels than may frequently be encountered on lichens. situated well north of the Antarctic Convergence and normal the wetting effect is prolonged as spray is The lower supra-littoral, characterized by the the shores enjoy ice-free conditions the whole year returned over the section just wetted. blackening lichen of the genus Verrucaria, is round. It is therefore somewhat surprising that all inhabited by large numbers of orange mites forms of littoral invertebrates are cryptic to a marked Biotic factors (Mesostigmata) and Halozetes marionensis which degree. Casual observation on any shore of these Habitat modification: In many instances tt was noted live in microdepressions of the lichen thalli. The islands will give the impression that there is no animal that certain organisms modify the habitat thereby siphonarid Kerguelenella lateralis o.ccurs at this level life at all, and it is often only after prolonged facilitating invasion by other forms. In the upper sub­ also and Knox96 (1968) has suggested that herbivor­ searching that certain groups may be detected. littoral of the Transvaal Cove Transect encrusting ous molluscs may well depend on Verrucaria as a All organisms living in the littoral are adapted to algae grow between and around adjacent small major source of food. Simpson97 ( 1976) however has withstand considerable inundation by fresh water boulders to form a uniting matrix which stabilizes the recorded only "green algal film" ( Ulothrix?) as a food since there is precipitation of some form on an substrate. Boulders here remain in one position for item of Kerguelenella on Macquarie Island. average of 25 days of every month. Certain organisms long enough to permit successive colonization by The fronds of Porphyra and Rhodymenia species I tend to favour fresh water and grow on faces down a variety of forms. A wide variety of invertebrates can provide refuge for large numbers of amp hi pods of the which fresh water seeps (e.g. Rama antarctica, live in safety beneath the boulders. genus Hyale, and to a lesser degree oligochaets Plate 11 . Cliff fragmentation results in large angular boulders Bostrychia spp. and Enteromorpha spp. at Duiker's Durvil/ea antarctica is effective in damping the Lumbricillus spp. Algae such as Bostrychia and forming a new shore line (Rook's Bay). Point Transect and at the small cove just north of the swell where it extends seawards on a gently sloping Rama are colonized by the minute pelecypod l..Asaea 32 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 33

encrusted, suggesting that no vertical migration locality to differ significantly from the sub-Antarctic occurs. Regular sampling over at least one year is pattern on the basis of algae. This is not surprising needed to resolve the movements of Nacella and to since no other island lies on or to the north of the establish whether there is more than one population. Subtropical Convergence as Tristan does. Del/108 (1972) has convincingly pointed out the dangers of assigning a particular locality to a Zonation in the Sub-Antarctic biogeographic region or province on the basis of common species in a certain group. The wide variety It is possible now for the first time to compare of reproductive methods, duration of larval phases zonation throughout the sub-Antarctic region. Knox99 summarized the zonation known by 1968 for and abilities to migrate pelagically or benthically five sub-Antarctic areas (Magellanic South America, encountered throughout benthic invertebrates does not permit the designation of generalized marine Falkland Islands, Gough Island, Auckland Islands, provinces where isolated islands separated by great Macquarie Island) and was able to draw up a scheme distances and abyssal depths are concerned. Thus it is representing a basic pattern common to all of them. interesting to note that the littoral fauna of Tristan To this list we can add Tristan da Cunha (since has strong affinities with Gough Island, Magellanic included in the sub-Antarctic by Dell, 1oo 1972), South America and the Falkland Islands (Chamber­ Marion and Prince Edward, Crozet, Kerguelen and lain, Holdgate & Wace, 109 in preparation) Camp bell. Following Knox'siOI (1968) basic scheme, zonation As expected, littoral zonation on Marion and may be represented as given in Table 4. Prince Edward corresponds closely with the basic It is at once clear that Tristan da Cunha is the only pattern, but more closely still with Crozet and

Table 4 Plate 13. Seals and penguins smother and abrade littoral communities on a small boulder beach at Macaroni Bay. Zonation on sub-Antarctic islands since Knox (1968)

Basic Pattern Tristan Marion and Crozet Kerguelen Campbell da Cunha Prince Edward consanguinea, by Lumbricil/us and insect pupae. The supra-littoral, upper and mid-littoral occur at Lichens Lichens Lichens Lichens Mixed turf carpets of Rhodymenia species 2 are inhabited by different times of the year. In November a band of Upper supra- Lichens littoral Caloplaca Prasiola Prasiola Lichens the mite Halozetes marinus devilliersi, Lasaea, Ulothrix sp. makes its appearance in the lower supra !.Re idea Lichina Kidderia, the oligochaets Lumbricillus macquariensis littoral where it persists through the summer and j:Juel/ia etc. and L. aestuum, and the polychaet Pionosy//is nurrix. dies off early in winter. The holdfasts of Durvillea support a varied fauna of Porphyra first becomes prominent in December Lower supra- Verrucaria Porphyra Verrucaria Verrucaria Verrucaria (Verrucaria) sponges, polychaets, amphipods, bryozoans, pelecy­ and grows steadily in the upper and mid-littoral, littoral Hildenbrandia Bostrychia Ulothrix Ulothrix Ulothrix Hildenbrandtia pods, gastropods and holothuroids. Isopods Dyna­ attaining luxuriance in mid-summer. It would appear Blue greens Halozetes Urospora Urospora menella and amphipods Hyale and Jassa crowd the from several observations that Rhodymenia species I Collembola fronds of red algae in the sub-littoral together with reaches its peak growing season in mid or late Diptera gastropods, pelecypods and copepods. Gaimardia summer. This would account for the abundance of Codium Rama Porphyra Porphyra Porphyra trapesina is abundant on Macrocystis fronds together Rhodymenia to the exclusion of all other algae in the Upper Porphyra littoral Bostrychia Bostrychia Acrosiphonia Diptera Bostrychia with various hydroids, and Lissarca rubrofusca is mid-littoral of Boulder Beach on the north-east coast Limpets Porphyra Coleoptera Algal carpet common on Desmarestia. of Prince Edward Island in April. Mention has been Siphonarids Kerguelenel/a Kerguelenella Birds and seals: At various points on the island made of the severe wave action at that point, and the La sa ea penguins and seals come ashore in huge numbers and frequently rotating boulders could only play host to abrade the substrate clear of organisms (Plate 13). organisms whose growth flush coincided with a Mid lridaea Gigartina Porphyra lridaea lridaea Rama Their influence is most marked between the upper period of stability. At Transvaal Cove individual littoral Limpets Mixed algal Porphyra Porphyra Kerguelenel/a supra-littoral and the mid-littoral, but once the lower boulders colonized by Porphyra on one side and Siphonarids turf Acrosiphonia Acrosiphonia Xiphophora Rhodymenia Mussels littoral has been reached they are borne up by water. Rhodymenia on the other were frequently encountered. Barnacles Rhodymenia Rhodymenia Kerguelenel/a Kerguelenella Large penguin colonies such as those found at Mixed algal Stable experimental rock faces stripped bare at Amphipods Bullard Beach and Kildalkey Bay make inroads into turf different times of the year and checked weekly for Mussels the maritime plant communities thereby wearing organisms would yield valuable information on down the vegetation and exposing the soils to growing seasons. Lower Dun'illea Lithothamnion Durvillea Durvil/ea Dun'il/ea Durvillea erosion. This together with abundant guano and In midwinter Porphyra seems to disappear entirely littoral Lithothamnion Halopteris Lithothamnia Lithothamnia Lithothamnia Lithothamnion feathers severely discolours the sea in the vicinity from the shore while scant growths of Rhodymenia Lithophyllum Laminaria Nacella Patinigera Camharidus after even light rainfall. The effect of restricted light persist (data from photographs). It is not known what Melobesia Hemiarthrum penetration in the upper sub-littoral as a result of this happens to the rich populations of amphipods at such is not known. It is likely too that eutrophication will times. Upper sub- I.Rssonia Macrocysris Po~l'siphonia Polysiphonia Po~rsiphonia I.Rssonia have considerable effect on benthic communities. Walker98 (1972) has shown that the Antarctic lim­ littoral Macrocyst is Schizoseris Schizoseris Schi;oseris Macrocyst is More obvious effects of guano deposition could be pet Patinigera polaris comprises two populations, Desmarestia Desmarestia /ridaea lridaea Macrocyst is Patinigera detected in the upper supra-littoral where Prasiola one of which migrates into the littoral from the sub­ Mixed red Nacel/a crispa grows luxuriantly below rocks favoured as bird littoral in summer. Nacella delesserti on Marion has algae an interesting distribution, juveniles being common perches. (Data from (Algae only !Data from own (Algae only (Data from Dele· (Data from Mor- in the kelp zone while vast carpets of adults occur in Knox. 102 1968) shown. Data from observation) shown. Data from pine & Hureau.I05 ton & Miller.'"' Season the sub-littoral. The shells of cryptic adults are Baardseth. 101 Delepine & 1966 and Ar- 1968) Marked changes in zonation patterns of the lower lithothamnion-free, while those in the open are 1942) Hureau.'o-c 1966) naud.'"" 1974) 34 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 35

Kerguelen, in that these groups lack limpets in the been identified to species level but all other specimens upper littoral and barnacles and indigenous muss.els belong to well-known species. in the mid-littoral. Macquarie Island is similar in this Of eleven species of Hydrozoa from the islands, respect. two were described as new by Millardll 7 (I 971 ). Only one species (Obelia geniculata) is cosmopolitan, the rest being widely distributed throughout the Antarc­ Universal schemes for littoral tic and sub-Antarctic. An interesting record from the zonation 1972/73 expedition was Halicrvstus antarctica (Scy­ phozoa: Stauromedusae), previously known only Littoral ecologists have long been concerned with from South Georgia and Grahamland (Millard, drawing up zonation schemes which would be personal communication). E applicable to patterns on nearly all seashores of the Recorded species of polychaetous worms now total E world. This was stimulated by the fact that certain 33, of which only one is positively new to science and lt) organisms or types of organisms tend to colonize the three show local variations from the norm. Seven same levels on shores geographically far apart. The species are cosmopolitan while the rest are of work of Stephenson & Stephenson"o (1949) was Antarctic and sub-Antarctic distribution. Day118 regarded as the most important, and is still used by ( 1971) found that one species ( Orbiniella minuta) was many workers in spite of criticisms and more recent known previously from Tristan da Cunha only. schemes. (e.g. Womersley & Edmonds,''' 1952; Hirudineans and oligochaets from the island's shores Lewis, 11 2 1972). Nearly all the schemes employ each comprise two species well known from high littorinids, barnacles and laminarians as indicators of southern latitudes. particular tidal levels. The only workers who appear Of seven species ·of isopods thus far collected, two to have attempted to correlate zonation on a sub­ were described as new by C/eretiJ9 (1971). Most of the Antarctic island (Macquarie) with the universal amphipods are typical southern forms and most of scheme are Kenny & HaysomliJ (1962), but they them have been recorded from Crozet and Kerguelen pointed out that this was not possible on account of as well. It is interesting to note that all three oribatid the total absence of littorinids and barnacles there. mites thus far collected from Marion's shores are new They suggested though that the Durvillea zone could taxa. However these animals are extremely tiny and be considered as the laminarian zone of the typical inconspicuous, and it is likely that in time these plan. 4 species will be found on other sub-Antarctic shores as The more recent scheme of Lewis" (1972) suggests well. that the littoral zone may be interpreted as falling Collections of echinoderms from Marion's shores between the upper limit of Littorina / Verrucaria and now comprise nine species, none of which are new to the upper limit of laminarians. The zone is further science since the new species Nul!amphiura marionen· divided into a littoral fringe and a eulittoral zone by sis (Bernasconi) has been synonymized with Amphiura the upper limit of barnacles. The upper limit of tomentosa (Lyman) by Rowe & C/arkei20 (1975). laminarians marks the top of the sub-littoral zone, The species are widespread throughout sub-Antarctic and this would tie in with the contention of Arnaud"5 and Antarctic shores. The algae comprise at least 60 (1974) that Durvillea is a sub-littoral organism. The species and show affinities with those of high only difficulty resides in the absence of barnacles or southern latitudes. However lack of reproductive F similar organisms in the mid-littoral to separate the structures on much of the material gathered posed littoral fringe from the eulittoral zone. taxonomic problems, and identification to species On Marion the upper limit of Porphyra would level of many genera awaits a future expedition. serve the purpose in summer, but this rhodophyte disappears in winter. It is not known with what Marion and Prince Edward have very few endemic permanence the orange mite (Mesostigmata) occupies species. At first glance this would seem to be an the shores, but it may form a useful indicator in the anomalous situation, since oceanic islands separated mid-littoral. The writer believes that the Lewisl66 from continents and other islands by barriers such as (1972) zonation scheme is an advance over other great distances and abyssal depths generally favour schemes because of its simplicity, its applicability to high rates of speciation. Another interesting feature the widest variety of conditions, and the fact that no of Marion and Prince Edward Island's littoral fauna is attempt is made to correlate organisms on the shore the low number of species, which together with their ![ with astronomical tidal heights. predominantly small size and cryptic nature give the island's shores an appearance of extreme impoverish­ ment. Reasons for these features may be gained from a Biogeographical notes consideration of the islands' histories, and compari­ Distribution records of invertebrates from the shores son with other islands of similar latitude. Van of Marion and Prince Edward indicate that the Zinderen Bakkerl21 (1971) has pointed out that the littoral affinities of these islands lie predominantly far greater age of islands such as the Crozet and with the sub-Antarctic region and to a lesser degree. Kerguelen archipelagos would allow not only coloni­ with the Antarctic. zation by more species but also a greater degree of The mollusc species are widely distributed south of speciation. It is likely too that extinction rates during the Subtropical Convergence from Magellanic South the glacial phases on Marion and Prince Edward were America around the world and to Antarctica. Three high, and that the present littoral fauna comprises the specimens of the genus Philobrya (Pelecypoda) Pla te 14. Common invertebrates from the shores of Marion Island. A: Kidderia sp. in Rho(~l'/1/enia turf: B. Lasaea wnsanguinea among few surviving species and recent migrants. Islands tangled fila ments of Bowrychia; C: Romanchella sp. aggregation on a stone from beneath boulders: D: J\'acel/a ( Patinigera)delesseni on a collected during the 1972/73 expedition have not such as the Crozet and Kerguelen groups did not shaded lithothamnia-free boulder; E: Kerguelenel/a lateralis in a Vermcaria·fillcd depression: F: Hemiarthrum seruloswn on lithothamnia. S. Af r. J. Antarct. Res., 1976, Suppl. 1 36 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 37

experience glaciation of the same magnitude as type of substrate is complex. Simplest conditions are REFERENCES Marion island and it is probable that a greater found where smooth very steep cliffs occur on the Order of Zinderen Bakker Sr., J .M. Winterbottom number of forms survived (van Zinderen Bakker:l22 relatively sheltered east coast. Here the swells merely appearance & R.A. Dyer, 384-390. Cape Town, A.A. 1971). Furthermore the relatively high number of rise and fall against the cliff face, and kelp is often the in text Balkema, 1971. 118 islands in those groups (particularly Kerguelen) could only visible organism. With increased exposure the Deacon, G.E.R. The hydrology of the have acted as reservoirs for prompt repopulation to Alston, A.H.G. & Schelpe, E. The Pterido­ development of a spray and splash zone results in the Southern Ocean. Discovery Reports, 15, nearby islands where fauna may have been annihilated frequent drenching of surfaces well above sea-level. phyta of Marion Island. Journal of 40, 60, by volcanic activity. 1-123, 1937. 63 Turf-like carpets of algae grow luxuriantly above the South African Botany, 23, 105,-109, 1957. 11 It is tempting to suggest that much of Marion's Doty, M.S. Rocky intertidal surfaces. Mem. kelp and form refuge for invertebrates. However the Andriashev, A.P. A general review of the shore fauna that"inhabits depressions and rock clefts geol. Soc. Am., 67(1), 535-585, 1957. 83 appearance of vertical cliffs suffering extreme Antarctic fish fauna. Biography and is a relict fauna from glacial times when sea Delepine, R. & Hureau, J.C. Comparative exposure is often similar to that of vertical cliffs with ecology in Antarctica. Monographiae temperatures were lower and ice scoured the coasts. minimal exposure. There thus appears to be a limit study of the vertical distribution of the bioi.. IS, 491-550, 1.965. 124 Littoral organisms in Antarctica show a strong beyond which increased exposure reduces the biota. marine vegetation of Archipel de Kergue­ Antarctic Pilot. Published by the Hydrogra­ preference for crevices. Conditions are complicated by movable substrates len and Iles Crozet. In Symposium on The Kerguelenian Province: On the basis of broad (e.g. boulders) which are common on Marion. phic Department under authority of the Antarctic oceanography, 164-165. Cam­ 31. 53, faunistic similarities workers in the past have devided Surface area available for colonization increases with Lords Commissioners of the Admiralty. bridge, Scott Polar Research Institute, London, 1961. 54. 104. the Antarctic and the sub-Antarctic region into decreasing boulder size. Also, the density of packing 6 1968. 105 provinces, districts and sub-regions (e.g. Knox, 123 increases with roundness of boulders. However, Arnaud, P.M. Contribution a la bionomie Delepine, R. La vegetation marine dans 1960; Andriashev, 124 1965; Kusakin, l25 1967; Hedg­ round boulders are indicative of frequent rotation due marine benthique des regions antarctiques l'Antarctique de l'ouest comparee a celle peth126 1969). Marion and Prince Edward islands to wave action, and no organisms can survive on et subantarctiques. Tethys, 6(3), 465- 33,92 des 'iles australes francaises. Consequences have generally been grouped together with the frequently abraded surfaces. The smaller the boulders 656, 1974. 106. 115 biogeographiques. C. r. somm. Seanc. Crozets, Kerguelen, Macquarie and sometimes Heard the more likely are they to be rotated. It is instructive Baardseth, E. The marine algae ofTristan da Soc. Biogeogr., 374, 52-68, 1966. 32 as the Kerguelenian Province or subregion. to note that boulders of approximately 70 cm Cunha. Results of the Norwegian Scienti­ Dell, R.K. Antarctic benthos. In Adv. Mar. 127 Del/ ( 1972) has convincingly expressed the severe diameter are well covered on calm shores but are fic Expedition to Tristan da Cunha, 1937- 20, 100, 28. 58, Bioi.. edited by F. Russell and M . Y onge, 108. 127. limitations of the provincial concept and has argued smooth and devoid of life on exposed shores. The 38, (9), 1-173, 1942. 81 . 103 10, 1-26. London, Academic Press, 1972. 130. 131 that a barrier "which will act on all organisms in a best protected and stablest niche is the substrate Ballantine, W.J. A biologically defined Dickie, G. Algar collected by Mr Moseley at similar way causing a large number of geographic beneath several layers of boulders. Accumulated exposure scale for the comparative descrip­ Marion Island in 40 fathoms. J. Linn. Soc. ranges to cease at about the same point" is needed detritus retains water at low tide and forms suitable tion of rocky shores. Field Studies. 3( I), I­ (Botany), 15, 42-43, 1877. 5 before a provincial boundary can be established. The material for in-fauna such as burrowers, tube builders I 9, 1961. 48 Engelbrecht, C.M. The genus Halozetes wide variety of modes of reproduction, duration of and under-stone dwellers. Bennett, I. Shores of Macquarie island. p. (Oribatei: Acari) on Marion Island. larval phases, dispersal ability and degrees of 69. Adelaide, Rigby Ltd., 1971. vulnerability that occur in the many groups of 36 Navors. nas. Mus. , Bloemfontein, 3(1 ), ACKNOWLEDGEMENTS 1974. invertebrates preclude comparisons from a common Bennetts, W.A. Island home of the wander­ 17 The author is indebted to the South African frame of reference, and McDowa/1128 (1968) has ing albatros. African Wildl(fe, 2(4), 22- Fuller, N. R. A preliminary report on the Department of Transport and SASCAR for finan­ 31, 1948. criticized Briggsl29 ( 1966) for his unrealistic compari­ 8 litoral ecology of Marion and Prince cial support, logistics and permission to carry out this 13.41 sons of this nature. Del/130 ( 1972) has further pointed Branch, G.M. The ecology of Patella. V. Edward Islands. S. A.f/". J. Sci.. 63(6). 248- investigation. Special thanks are due to Professor 49. 51, out that littoral distribution patterns may not Commensalism. Zoologica A.fricana, 10, 252. 1967. 65 E.M. van Zinderem Bakker for accommodating this correspond with those on the shelf or in deeper water, 133-162, 1975. 75 Hedgpeth, J . W. Marine biogeography of the project in his overall programme. and that provincial comparisons should be limited to Briggs, J .C. Oceanic islands, endemism and Antarctic regions. In Antarctic ecology, The following specialists, identified material and specific groups of organisms living between certain marine paleotemperatures. Syst. Zoo/. 15, edited by M. W. Holdgate, 97-104. Lon­ are gratefully acknowledged: depths. Dell concludes that to state that an organism 153-163, 1966. don, Academic Press, 1970. 126 Mrs F. Arnaud, Pycnogonida; Dr P. Arnaud, 129 occurs in the Kerguelenian province does not tell one Chamberlain, Y. M. Marine algae of Gough Huntley, B. Vegetation. In Marion and very much. It may occur on Kerguelen or Macquarie Mollusca; Dr A. Clarke and Dr S. Rowe, Echinoderms; Miss P. Cook, Bryozoa; Dr E. Easton, Island. Bull. Brit. Mus. Nat. Hist. (E: Prince Edward Islands. Reports on the or both. It may also occur off South Georgia, in I 0 to Oligochaets; Dr C. Engelbrecht, Oribatidae; Dr B. Botany) 3(5), 175-232, 1965. 29. 89 South African Biological and Geological 75 metres and in the Ross Sea in 300 metres. More Kensley, Isopoda and Tanaidacea; Dr P. Knight­ Chamberlain, Y. M., Holdgate, M. & Wace, Expeditions, 1965-66, edited by E.M. van useful .information would be that a species has been Jones, Spirorbinae; Dr V. Koltun, Porifera; Dr D. N. The littoral ecology of Gough Island, Zinderen Bakker Sr., J.M. Winterbottom recorded from Kerguelen in 10 to 75 metres and in the 30, 55, South Atlantic. (In preparation). 90. 109 & R.A. Dyer, 98-160. Cape Town, A.A. Ross Sea in 300 metres. Lindsay, Mycophyta, Dr R. Miller, Ascideacea; Or N. Millard, Hydrozoa; Dr G. Murina, Sipunculoidea; Balkema, 1971. 67 The present writer agrees entirely with the views Cleret, J.J. Isopoda. In Marion and Prince Dr N. Riser, Nemertea; Dr R. Sawyer, Hirudinea; Mr Jeannel, R. Au seuil de I'Antarctique. expressed by De/11 31 as far as marine littoral Edward Islands. Report on the South R. Simons, Phycophyta; Mr V. Smith, Bryophyta. Croisiere du ·· Bougaim'ille" aux 'iles des organisms are concerned. African Biological and Geological Expedi­ Much assistance was received before and after the tions. 1965-66. edited by E. M. van manchots et des elephants de mer. Paris, expedition. Thanks are due to Mr G. Altham, Mr R. Zinderen Bakker Sr., J.M. Winterbottom Editions du Musee, 1941. 7 Butlin, Mr A Crawford, Mrs A. Gerber, Mr A.M. & R.A. Dyer, 379-383. Cape Town, A.A. Kenny, R. & Haysom, N. Ecology of rocky General conclusions 34, 52, Shipley, Miss B. Southey and Commander C.J.H. Balkema, 1971. 119 shore organisms at Macquarie Island. 70, 76. The generally low density of organisms and low Wagenfeld. Crawford, A. B. The birds of Marion Island, Pacif Sci .. 16(3), 245-263, I 962. 91. 113 number of species indicate the instability of Marion's The author is indebted to Or P. Arnaud for South Indian Ocean. Emu. 52(2), 73-85, Kensley, B. Five species of Jaeropsis from littoral environment. In addition to sparse coverage critically reading the manuscript and for helpful 1952. 9 the southern Indian Ocean (Crustacea, within niches, there are also apparently many vacant suggestions, and Dr J. Grindley for advice in certain Isopoda: Asselota). Ann. S. A.fr. Mus .. niches. The drastic effects of glacial and volcanic Day, J.H. The biology of Langebaan problems and comments on the manuscript. 67( 10), 367-380, 1975. 19 activity within relatively recent times ( 100 000-15 000 Lagoon. A study of the effect of shelter Finally thanks are due to those who helped during Knox. G.A. Littoral ecology and biogeogra­ years B.P.) are not the only causal factors involved. from wave action. Trans. Roy. Soc. S. 21, 42, expeditions: Mr D. Gerneke who participated in all A.fr., 35(5), 475-547, 1959. phy of the Southern Oceans. Proc. R. Soc. Large swells and exposed or unstable substrates also the field work, and Dr J. Grobbelaar, Mr P. Condy, 43 69. 72, Day, J.H. Polychaeta. In Marion and Prince (B), 152, 577-625, 1960. 87. 123 render the littoral environment hostile to life. Mr L. Heinonen, Mr J. Langford, Mr F. Potgieter Edward Islands. Report on the South The interplay between intensity of wave action and and Mr H. Thomson. Knox, G.A. The biogeography and intertidal African Biological and Geological Expedi­ ecology of the Australasian coasts. In tions. 1965-66. edited by E. M. van Oceanography and marine biology. Annual 38 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1 S. Afr. J. Antarct. Res., 1976, Suppl. 1 39

review. edited by L. Bames, 1, 341-404. Rowe. F.W.E. & Clarke, A.M. Notes on APPENDIX London, Alien & Unwin, 1963. 22,37 some echinoderms from Marion Island. Knox, G.A. Tides and intertidal zones. In 23, 45. Bull. Br. Mus. not. Hist.(D: Zoology), Species lists Symposium on Antarctic oceanography, 73, 74, 28(5), 187-190, 1975. 18, 120 78. 88. 131-146. Cambridge, Scott Polar Research 96. 99. Sawyer, R.T. A new species of "tentacled" Angiosperms, lichens. bryophyte~. fungi and algae were recorded from the following habitat levels: Institute, 1968. 101. 102 marine fish leech parasitic on Notothenia I upper supra-littoral Knox, G.A. Antarctic· marine ecosystems. from the Subantarctic Marion and Crozet 11 lower supra-littoral In Antarctic ecology, edited by M. W. Islands. Hydrobiologia, 40(3), 345-354, Ill upper-littoral Holdgate, 1, 69-96 London, Academic 1972. 16 IV mid-littoral Press, 1970. 24.62 Schul7e, B.R. The climate of Marion Island. V lower littoral Kusakin, O.G. Fauna of Isopoda and ln Marion and Prince Edward Islands. VI upper sub-littoral, lower red zone I Tanaidacea in the coastal zones of the Report on the South African Biological VII upper sub-littoral, lower red zone 11 Antarctic and Subantarctic waters. lssled. and Geological Expeditions, 1965-66, Fauny morei.. 4( 12), 320-389, 1967 (in edited by E. M. van Zinderen Bakker Sr., Invertebrate habitats were further defined as: Russian). 125 J.M. Winterbottom & R.A. Dyer, 16-31. 39. 61, A on exposed rocky surfaces La Grange, J.J. Notes on the birds and Cape Town, A.A. Balkema, 1971. 66,95 B partially sheltered (on sides, overhangs, crevices or depressions) mammals on Marion Island and Antarc­ Simpson, R. Physical and biotic factors C more or less completely sheltered (under-surfaces, substrates or pools) tica. Jl. S. Afr. bioi. Soc., 3, 27-84, 1962. 12 limiting the distribution and abundance of D on or among fronds and under holdfasts of algae littoral molluscs on Macquarie Island Langenegger, 0. & Verwoerd, W.J. Top­ 35. 77. ographic survey. In Marion and Prince (Sub-antarctic). J. exp. mar. Bioi. Ecol., 79, 80. ANGIOSPERMAE Lithoph_rllum sp. VI. VII Edward Islands. Report on the South 21, 11-49, 1976. 94,97 Corula plumosa Lophurella sp. V. VI, VII Skottsberg, C.J.F. Communities of marine Tillaea moschata L. hookeriana VI. VII African Biological and Geological Expedi­ Macrocystis p_rrifera edited by E.M. van algae in Subantarctic and Antarctic VII tions, 1965-66. BRYOPHYTA ,tf_rriogramme s p. VI, VII Zinderen Bakker Sr., J.M. Winterbottom waters. K. Svenska Verensk.Akad. Hand/., 25, 56. Eriopus sp. Neuroglossum sp. 19(3), 1-92, 1941. 57 VI & R.A. Dyer, 32-39. Cape Town, A.A. 50 Dicranoll'eisia sp. Ph_rmatolithon sp. VII Balkema, 1971. 47. 68, Stephenson, T.A. & Stephenson, A. Univer­ Phyllophora appendiculata IV Lewis, J. The ecology of rocky shores. 82. 86, sal features of zonation between tide MYCOPHYTA Po~rsiphonia sp. VI. VII 93. 112, Lichens P. urbanoides London, English Universities' Press, 1972. 114.116 marks on rocky shores. J. Ecol., 38, 289- 59. 84, VII Buellia coniops I Porph_rra sp. Ill. IV McDougall, I. Geochronology. In Marion 305. 1949. 110 Stephenson, T.A. & Stephenson, A. Life Caloplaca cirrochrooides I Porph_rropsis l'exil!aris V and Prince Edward Islands. Report on the Haematomma er_rthromma I Prasio/a crispa I. If South African Biological and Geological between tide marks on rockr shores. San Francisco, W.H. Freeman, 1972. Lecidea interrupra I Rama antarctica 11. Ill, IV Expeditions, 1965-66, edited by E.M. van 27 Pertusaria sp. I Rhi:oc/onium riparium Straehl, A. A. Aspectos ecologicos de algunas 11. Ill Zinderen Bakker Sr., J.M. Winterbottom Verrucaria resselatula 11 Rhodochorlon sp. I. 11 & R.A. Dyer, Cape Town, A.A. Balkema, areas intercotidales de la costa chilena V. durietzii I Rhodoglossum spp. IV, V, VI, VII 197 1. 2 entre Ios paraleles 36° y 54° lat. Su Univ. V. mucosa 1-V Rhodophyllis sp. VI Concepcion, Chile, Inst. central de Biolo­ V. maw·a I Rhoc~rmenia spp. McDowall, R.M. Oceanic islands and Ill, IV, V Fungi Schi:oseris sp. endemism. Syst. Zoo/., 17(3), 346-350, gia, Dept. Zoo! (unpublished thesis) I-XI, VI. VII 1- 114, 1964. 26 Discothecium gemm(/'erum Sphace/aria sp. VI 1968. 128 Verwoerd, W.J. Geology. In Marion and Spon!(omorpha arcta VI Millard, N.A. Hydrozoa. In Marion and PHYCOPHYTA Spon!(omorpha pacifica VI, VII Prince Edward Islands. Report on the Prince Edward Island~. Report on the A broteia suborbicularis IV. V Ulothri.\' sp 11, Ill Sourh African Biological and Geological Arcrosorium sp. V. VI U/1'0 sp. VII South African Biological and Geological Expeditions, 1965-66, edited by E.M. van Expeditions, 1965-66 edited by E.M. van Acrochaetium sp. I. 11 Zinderen Bakker Sr., J.M. Winterbottom A denoc_rstis utricularis VII PORIFERA Zinderen Bakker Sr., J.M. Winterbottom & R.A. Dyer, 40-62. Cape Town. A.A. Ballia callitricha VI. VII Amphi/ecws fucorum VI. B. C & R.A. Dyer, 396-408. Cape Town. A.A. I. 44, Balkema, 1971. 46 Bostr_rchia mixta I I. Ill Clarhrina coriacea VI, C Balkema, 1971. 117 B. 1·aga 11, Ill Ect_rom_rxilla chilensis VI. B. C Mitchell-Innes, B.A. Primary production Walker, A.J.M. Introduction to the ecology B. typica Ill Halichondria panicea VI. B. D studies in the south-west Indian Ocean, of the Antarctic limpet Parinigera polaris Ceramium spp. VII Haliclona topsenri VI. B. C 1961-1963. Oceanographic Research Insti­ at Signy Island, South Orkney Islands. Br. Ceramium rubrum VII Ha/isarca dujardini VI. B Cladophora spp. tute lnvesrigational Report, 14, 1-20, 1967. 14 Amam. Sun·. Bull.. (28). 49-69. 1972. 98 11. Ill Leuconia joubini VI. B. C Cladophora subsimplex 11. Ill Leucosolinia borryoides VI. B Morton, J. & Miller, M. The New Zealand Womersley. H.B. & Edmunds, S.J. Marine Codium sp. VI S.rcon kerguelensis VI, B. C sea shore. London, Collins, J968. 38. 107 coastal zonation in Southern Australia in Colacodas_ra inconspicua VI Moseley, H.N. Notes by a naturalist. relation to a general scheme of classifica­ Corallina spp. VI, VII BRACHIOPODA London, Murray, 1892. 4 tion. J. £col.. 40, 84-90. 1952. 85. Ill Cr_rptopleura sp. VII I specimen (unidentified) VI. C Delesseria sp. V. VI Ne!, E.A. The microplankton of the south­ Van Zinderen Bakker, E.M. Sr. Introduc­ Desmarestia sp. VI CNIDARIA west Indian Ocean. Invest!. Rep. Div. Sea tion. In Marion and Prince Edward D. chorda/is VI. VII Fish. S. Afr., 62, 1-40, 1968. 15 Hydrozoa Islands. Report on the South African D. rossii VI. VII H_rdractinia parvispina VI. B Orren, M .J. Hydrology of the south-west Biolof?ical and Geological Expeditions. Dun•il/ea?antarctica V. VI. VII (infertile thus provisional Indian Ocean. Invest/ Rep. Div. Sea Fish. 1965-66. edited by E.M. van Zinderen Ectocarpus spp. 11. Ill. IV. V. VI identification only) S. Afr., 55, 1-19, 1966. 64 Bakker Sr., J.M. Winterbottom & R.A. Enteromorpha bulbosa Il l. IV Geminocarpus geminatus VII VI. VII. B. C. D Rand, R.W. Notes on the birds of Marion Dyer, 1-15. Cape Town, A.A. Balkema, Myriothe/a meridiana Halopteris funicu/aris VI. VII Obelia genicu/ara VII, D Island. Ibis., 96(2), 173-206. 1954. 10 1971 . 121, 122 Heterosiphonia sp. VI, VII Sertularel/a picta VI, VII. A. B. D H. merenia VI. VII SympleC1osc_1phus marionensis VII, A, B Leptosomia simplex V Tu bulariidae VII, B 40 S. Afr. Tydskr. Antarkt. Nav., 1976, Bylae 1

Scyphozoa Kerguelenella lateralis II, Ill, IV, V, B Halic/ysrus antarcrica VII. D Nacel/a ( Patinigera) delesserri V, VI. VII, A, B Aanwysings vir Bydraers Actiniaria (English - see inside front cover) 2-3 species (not identified) VI, VII, B. C ARTHROPODA Crustacea en datum van publikasie. PLATYHELMINTHES Die doe! met die tydskrif is die verspreiding van inlig­ Amphipoda D ting oor Suid-Afrikaanse Antarktiese navorsing en bv. Roederer, J.G. Dynamics of geomagnetica//y Not identified V. VI. C, Cerapus tubularis VII. D verwante bedrywighede. Bydraes (in Engels of in trapped radiation. Berlin, Springer-Verlag. 1970. Hyale grandicornis Ill, IV. C. D Tydskrifartikels: Outeur(s), titel van artikel, volledige NEMERTEA H. hirtipalma V, VI, VII. C. D Afrikaans) wat nog nie voorheen gepubliseer of reeds Tetrasremma sp. VI. VII. C titel van tydskrif, volumenommer. eerste en Jaaste Jassa .falcata VI, VII, B. D elders vir publikasie aanvaar is nic, word gepubliseer VI. C in die volgende kategoriee. bladsy, datum. Pontogenia sp. bv. Rikitake. T. Electromagnetic induction in a ANNELID A lsopoda l. Navorsingsmededelinge - oorspronklike weten­ Polychaeta Dynamenella hwtoni VI, VII, C, D perfectly conducting plate with a circular hole. C. D skaplike verhandelinge. Boccardia polybranchia V, VI, Ewsphaeroma gigas VI, C Journal of Geomagnetism and Geoelectricity, V. VL VII. D 2. Kort berigte - die voorlopige mededeling van Cirratulus cirrarus !ais pubescens Commensal on Kroro. 16, 31-36. 1964. VI. VII, C. D oorspronklike navorsingsresultate. gegewens ens. Cirriformia sp. Exosphaeroma Die verwysing in die teks moet bestaan uit die naam Eulalia magalaensis VI, VII. B. C VII, D 3. Briewe aan die Redakteur - kort oorspronklike VII. B Jaeropsis curvicornis (name) van die outeur(s). gevolg deur die datum van Haploscolopos sp. Munna sp. V. D wetenskaplike mededelinge. Harmorhoe bergsrroemi V, VI. B, C. D 4. Oorsigte - van navorsing in verskeie dissiplines. die publikasie. Waar daar 'n groot aantal verwysings Copepoda Helicosiphon platyspira V, VI. B, C 5. Notas en nuus - kort verslae oor huidige bedry­ (gestel meer as 20) gegee word. moet hulle ook in die VI, VU, C, D Tigriopus angulatus IlL IV, C Nicolea venustula wighede, deelname aan Suid-Afrikaanse Antark­ teks opeenvolgend van opskrifnommers voorsien Oriopsis limbata V, VI, B, C, D Tanaidacea tiese ekspedisies, boekbesprekings ens. word, aldus: Paralaeospira /evinseni V, VI, B Anatanais gracilis IV, V, VI, VII. C, D Baker & GledhiiJ2 ( 1964) Phyllodoce polyphylla VII,C Die tydskrif word jaarliks gepubliseer. Navorsings­ Verwysings verskyn dan alfabeties volgens die naam· Pi/eo/aria tegwyni (Pro- VI, C mededelinge wat spoedige publikasie regverdig, kan Acarina van die eerste outeur in die lys aangegee, gevolg deur visional identification) egter onmiddellik na aanvaarding deur die redaksie V, VI, C, D Oribatidae die toepaslike opskrifnommers in 'n kolom aan die Pionosyllis nutrix as separate op koste van die betrokke outeurs (teen regterkant. Sodoende kan enige verwysing dan weer Platynereis australis V, VI, VII, D Halozetes marionensis ll. A. B Ill, IV. D die gewone oordruktariewe) uitgegee word. Po~rcirrus hamilroni VI, B, C H. marinus devil/ie~·si in die teks opgespoor word. P. kerguelensis VI, VII. B Mesostigmata Voorleggings van bydraes: Bydraes moet voorgele Tabelle moet op afsonderlike velle aan die einde van Potamilla antarctica VII. B, C 1-2 species 1ll. IV, A. B word in triplikaat (die oorspronklike plus twee die teks verskyn. Die nommer en opskrif moet bo aan Romanchel/a invenris V, VI, B, C afskrifte, volledig met tabelle, syfers ens) en dubbel elke tabel verskyn. VU, D gespasieer, slegs aan een kant van die bladsy en met Romanchella perrieri Pycnogonida lllustrasies: Die oorspronklikes moet ongeveer twee Syllis variegata VI. VII, C, D Tanystylum cavidorsum VI, VII, C, D ruim kantlyne getik. Elke bladsy moet 'n verkorting Tharyx sp VII,C van die titel en die skrywer se van daarop he. Referate keer so groot as die uiteindelike gedrukte groottes wees om vir fotografiese verkleining toe te laat. Die Thelepus setosus Vll,C, D BR YOZOA (provisional identification) kan Of in Engels Of in Afrikaans geskryf word. outeur se naam en die nommer van die figuur of plaat Crisia sp. VII, D Titel: Die titel moet kort wees (verkieslik nie meer as Hirudinea VII, D moet op elke illustrasie aangegee word. Letterwerk VII, D Hippothoa (Celleporella) hyalina tien woorde nie), maar moet insiggewend genoeg Antarcticobdella crozetensis Tubilipora spp. VII,D moet nie op 'n oorspronklike illustrasie aangebring VI, C wees vir gebruik in titellyste of by kodering vir die Glyptonotobdella antarctica Menipea crispa VII, B word nie, maar we! op 'n kopie wat die oorsponklike doe! van opberging en herkryging van inligting. Die vergesel, sodat eenvormige letterwerk van die vereiste Oligochaeta ECHINODERMATA outeur(s) se naam (name) en voorletters, affiliasies en grootte deurgaans aangebring kan word. Lumbricillus aestuum I!, Ill, IV, D huidige adres moet ook verstrek word. Lyntekeninge, insluitende grafieke, moet op teken­ L macquariensis IV, D Holothuroidea Pseudocnus laevigarus Vl, Vll. B, C. D Outeursamevatting: Dit kan tot 200 woorde lank wees linne, kunspapier van goeie kwaliteit of ruitjiespapier Sipunculoidea en moet die inhoud van die referaat feitelik en saaklik met blou lyne geteken word. Gebruik slegs werklik Golfingia ohlini VI, VII, B, C Ophiuroidea opsom. Dit moet in albei landstale geskryf word en swart lndiese ink. Lyne moet reelmatig wees en nie so Amphiura romentosa VII,C, D moet geskik wees vir afsonderlike publikasie en vir naby mekaar dat hulle in die drukproses ineenvloci VII, D MOLLUSCA Ophiurolepis martensi indeksering. nie. In grafieke moet krommes dik ker as aslyne Ophiacantha vivipara VII,D Amphineura Die hoofteks moet noukeurig en duidelik onder geteken word. Indien foto's verskaf word, moet dit VI, VII, C Lepidopleurus kerguelensis onderwerpshoofde uiteengesit word. Numerering en duidelike swart-wit-glansafdrukke met skerp kontra­ Hemiarthrum setulosum V. VI, VII, A. B, C Asteroid ea Anasterias rupicola Vl, VU. B. C voetnote moet vermy word en omslagtige aOeidings stering en detail wees. 'n Aanduiding van die ens. moet in die aanhangsels verskyn. vergroting moet deur middel van 'n skaal op 'n kopie Pelecypoda VII, D CHORDATA Korrekte spasiering sal vergelykings en formules in van die foto aangetoon word. Gaimardia trapesina Byskrifte wat in ooreenstemming met hul figure, Kidderia bicolor IV, V, VL C. D Ascideacea die teks laat uitstaan. Gebruik die solidus. met die ens. genommer is. moet afsonderlik getik en aan die K. oblonga V, VI. C. D Core/la eumyota VII, C nodige parentese, vir eenvoudige breuke. As koppe­ Lasaea consanguinea Il, III, IV, V, A, B, C Molgu/a sp. VI, B Iings en hakies nodig is. moet opgeboude breuke einde van die teks aangeheg word. Lissarca rubro.fusca VII, D Polr:oa rericu/ara VII, C gebruik word. Identifiseer handgeskrewe simbole in Redigering: Referate wat vir publikasie aanvaar Philobrya (3 species) IV, V, VL D S.nioicum sp. VI, B die kantlyn en verklaar alle simbole en veranderlikes word, sal ind ien nodig ter wille van korrekte of Styelidae. l:!otryllinae VII, C in die teks onmiddellik na die vergelyking. Die duidelike uiteensetting onderworpe wees aan redak­ nommer van 'n vergelyking kom tussen hakies en in teursveranderings, maar outeurs is verantwoordelik Pisces lyn met die vergelyking teen die regterkantse kantlyn. vir die feitelike korrektheid. Gastropoda Notothenia coriiceps Eatoniella kerguelenensis VI, C. D Notothenia macrocephala bv. (2) Proewe: Slegs een stel proewe word aan outeurs Laevilitorina caliginosa VL VII. C. D Harpagi.fer bispinis Metricke eenhede volgens die SI moet deurgaans gestuur en moet binne twee weke terugbesorg word. gebruik word. Publikasies van die Suid-Afrikaanse Verbeterings moet beperk word tot die korrigeer van Buro vir Stand<1arde oor die metrieke stelsel kan setfoute. Wysigings van die teks veroorsaak probleme hiervoor geraadpleeg word. en word as outeurskorreksies in rekening gebring. Verwysings: Bibliografiese verwysing moet die vol­ Oordrukke: Bestellings vir oordrukke moet by die gende inligting insluit: terugbesorging van proewe geplaas word. Vyftig oor­ Boeke: Out~ur( s ), titel. plek van publikasie, uitgewer drukke van elke hydrae word gratis verskaf.