Ecology of Rocky Shore Organisms at

RON KENNyl and NOEL HAYSOM2

IN 1948 the Antarctic Division, Australian De­ series of sand and shingle beaches separated by partment of External Affairs, established a re­ rocky headlands or reefs (Figs. 2, 3, 4) but in search station at Macquarie Island. As part of many places steep cliffs descend almost vertically the biological work a study of the rocky shore to the sea (Fig. 5). Much of the west coast has ecology of the island was made (by R. K. during extensive flat reefs and gradually sloping beaches 1948-49 and N. H. in 1949-50). often strewn with large boulders. Although large general collections have been The sites investigated in detail (Fig. 6) were made at Macquarie and other is­ all at the north end of the island and were se­ lands by various expeditions (notably the Aus­ lected to give a variety of habitats and exposures tralian Antarctic Expedition, 1911-13, and the to wave action. General observations were made British, Australian, and New Zealand Antarctic at other locations. Expedition, 1929-31) comparable ecological The climate is typically subantarctic. Meteoro­ programmes in southern latitudes have been logical details made available by the Common­ few. Of the papers on similar topics from tem­ wealth Meteorological Bureau, Melbourne, are perate southern latitudes those of Isaac (1937) given in Table 1. The air temperature range is on South Africa, Guiler (1952) on , small, the maximum recorded during the period Knox (1953) and Batham (1958) on South Is­ of this survey being 10.7 C (Jan 1950) and the land of New Zealand, and Guiler (19'59) on minimum -8.3 C (Aug 1950). Although the Chile are relevant to this study. total rainfall is not excessive, precipitation­ rain, hail, sleet, or snow-occurs almost daily. MACQUARIE ISLAND Strong winds, including gusts of more than 100 knots, are a striking feature of the climate and 0 0 Macquarie Island (54 29' S, 158 58' E) is are predominantly (more than 70%) from the situated midway between Tasmania and the An­ north and northwest. However, climatically the tarctic mainland. The island, with associated island is milder than other subantarctit;.. islands rock Outcrops to the north and south, is the ex­ of similar latitude and there is no permanent ice posed cap of an extensive submarine ridge run­ sheet. ning in a general north-south direction. The Heavy Seas occur at all seasons of the year, nearest neighbouring islands are the Auckland Is. and the coastline would be considered an ex­ and Campbell I., some 400 miles to the northeast posed one by any standard (Fig. 7). (Fig. 1) and separated from the Macquarie Macquarie I. lies within the area of Antarctic Ridge by depths of more than 2,000 fathoms. surface waters during part of the year, at least. The upper levels of this ridge are narrow, a The R.R.S. "Discovery II" plotted the Antarctic sounding of 1,548 fathoms having been recorded convergence in latitudes 53 0 and 54 0 S in this 5 miles east of Macquarie I. (Mawson, 1943). region during the winter months of 1932 A complete account of the geography and (Mackintosh, 1946). In the 1948-50 period the geology of Macquarie can be found in Mawson monthly mean sea temperatures ranged from (1943). The island, 21 miles long and up to 3 7.2 C (Jan) to 2.8 C (July). Surface salinities to miles wide, is largely composed of volcanic rock. the west and east of the island were recorded Most of the eastern coastline is formed of a by R.R.S. "Discovery II" during the 1932 winter and varied from 33.8%0 to 34.4%0 (Deacon, 1 Zoology Department, University of Queensland, 1937). Brisbane, . Manuscripr received June 1, 1961. 2 Fisheries Section, Department of Harbours and A tide gauge was set up in Buckles Bay (see Marine, Brisbane, Australia. Fig. 6) but was wrecked by storms after a short

245 246 PACIFIC SCIENCE, Vol. XVI, July 1962

150' E

__---+_ 50'S

• -Macquarie I.

__---I-60"S

120' E Southern Orran

___-_t___ 70' S

Ross Sea

FIG. 1. Macquarie Island and its relationship to surrounding land masses.

period of operation. From the records available (i) LICHEN ZONE; Extending from the area the spring range is 4 feet 6 inches and the neap dominated by maritime terrestrial flora range a little more than 2 feet. (Taylor, 1955), down the beach into the region of splash. The common organisms General Features of Zonation are lichens interspersed with some Six zones can be recognized on the rocky mosses. (ii) Porphyra ZONE: Porphyra umbilicalis shore extending from above high-water springs dominant from HWS3 to below HWN. to below low-water springs. The zones have (iii) "BARE" ZONE: Extending from just be­ been described in terms of "effective tidal low HWN to approximately MSL in heights" by Endean, Kenny, and Stephenson which the most obvious species was <1956) and other authors. These effective tidal Siphonaria lateralis. heights are based on the correlation of data --- 3 In this paper the following abbreviations have from the tide gauge at station 2 (see Fig. 6) been used: and from general observations at this and other high water spring tide, HWS stations. high water neap tide, HWN mean sea level, MSL The sequence of dominant organisms forming low water neap tide, LWN the zonation pattern is: low water spring tide, LWS Macquarie Island-KENNY and HAYSOM 247

TABLE 1 The faunal element of this zone is restricted METEOROLOGICAL DATA FOR MACQUARIE 1., to small species of cryptic habit living among 1948-50 the plants. The variable densities of these spe­ 1948 cies at different localities appear to be controlled Apr.- 1949 1950 by the amount of detritus, forming a suitable Dec. habitat, in the rock crevices and around the Air temp. °C plants. maximum...... ------. 8.75 9.65 10.7 The most frequently recorded animals are the minimum...... -3.65 -5.9 -8.3 mean...... - 4.25 4.35 mite Halozates sp., the beetles Antarctotachinus Total precip. (inches) ..... - 37.06 42.55 crozetensis and Antarctophytosus macquariensis, No. of days precip...... - 336 333 larvae of dipterans, Coleopa macquariensis and Mean wind vel. (knots).. - 18.2 19.3 - others, and various collembolans. Porphyra ZONE: Porphyra umbilicalis forms (iv) UPPER RED ZONE: In which Rhodymenia a thick mat, the upper limit of which is reached sp. is dominant from MSL to LWN. by the sea only at high water of spring tides. In (v) KELP ZONE: Durvillea stipes calm weather the dry weed forms a conspicuous cover almost the whole area of rock be­ band around the island's coastline. tween LWN and LWS, overlying a layer The upper parr of the zone is fo.rmed fre­ of encrusting red coralline algae. quently of an association in which as well as the (vi) LOWER RED ZONE: The red corallines dominant Porphyra there are many plants of extend below LWS and support a variety Rhizoclonium sp. and Iridea boryana and, less of red algae (Dellesseria spp., and Iridea commonly, Prasiola sp. In some situations Acro­ sp.) and brown algae (Desmarestia sp.). siphonia pacifica is found in patches, and this The most striking feamres of this zonation alga and Chaetangium fastigium are common in pattern are the absence of an upper "littorinid" the lower part of the Porphyra complex. and a "barnacle" zone, and the plant domination Where pools or similar suitable habitats are of the shore. In five of the zones the obvious available Viva lactuca, Enteromorpha intesti­ and dominant organisms are botanical, and al­ nalis, and Cladophora sp. extend throughout the though animals are common they are, generally, Porphyra mat. small and secondary to the algae and lichens in The animal species common at this level are biomass. It is only in the "bare" zone, where those noted in the lichen zone, with fewer Col­ Siphonaria lateralis is the commonest organism, lembola, and the addition of the oligochaetes that there is a faunal dominant. Lumbricillus macquariensis and Marionina anti­ podum. The gastropod Macquariella hamiltoni Details of Particular Zones has been observed in the Porphyra zone but LICHEN ZONE: This area is characterised by typically is found at lower levels of the beach. several species of lichen, the most obvious being "BARE" ZONE: Considerable variation in the Verrucaria sp. and an unidentified bright yellow horizontal and vertical extent of this zone was form, and the red alga, Hildenbrantia sp. observed (see below) and at some stations the Many plants, common in the terrestrial mari­ "bare" zone was not noted. However, when well time association (Taylor, 1955) extend seawards developed, it is marked by the close-cropped into this zone in varying densities. The moss nature of the algal constituents, the common Muelleriella crassifolia, and the grass Puccinellia forms being Chaetangium fastigium and Acro­ macquariensis, are more noticeable in relatively siphonia pacifica. exposed locations. The vascular plants Coloban­ The dominant organism is Siphonaria lateralis thus muscoides and Cotula plumosa, and the which, though common throughout this level of moss Ceratodon purpureus occur commonly in the beach, appears to favour dissected and crev­ more sheltered areas. Isolated plants of tussock iced rock surfaces. The species attains its maxi­ grass, Poa foliosa, are scattered through the up­ mum density at the lower limit of this zone per section of this zone. (approximately MSL) and in suitable locations 248 PACIFIC SCIENCE, Vol. XVI, July 1962 numbers up to 1,500 individuals per square plants of the dominant alga were observed metre. among the stipes of the kelp and extending be­ Other common animals include Macquariella low LWS. The dense growth of Rhodomenia hamiltoni, the limpet, Nacella delesserti, Lum­ forms a thick covering, protecting a wide range bricillus macquariensis, Marionina antipodum, of small animals. numerous amphipods, Hyale novae-zealandiae The fauna shows elements from the higher being the most frequently recorded, and many zones and also many species which are usually small nematodes. found at lower levels. Some specimens of Halozates sp. and Coleopa Hyale novae-zealandiae is, numerically, the macquariensis range down to this level but they dominant animal and Siphonaria lateralis, Na­ are essentially fauna of the upper zones. Simi­ cella delesserti, and Macquariella hamiltoni are larly a few nemerteans and turbellarians have the common secondary animals. Lumbricillus been recorded from the "bare" zone but they are macquariensis and Marionina antipodum were more common below MSL. recorded in small numbers. UPPER RED ZONE: This is a narrow zone in Other animals noted are representatives of which the red alga Rhodomenia sp. is the domi­ the fauna of lower tidal levels and are not as nant organism. In general, the boundaries of the common as those mentioned in the preceding zone are sharply defined. On certain reefs, how­ paragraph. They include the lamellibranchs ever, the lower limit was ill marked and many Kidderia pusilla and Gaimardia trapesina, the

FIG. 2. View from Wireless Hill looking south along rhe isthmus with Hasselborough Bay on the right. Station 4 in the right middle distance. (Australian Department of Information photo.) Macquarie Island-KENNY and HAYSOM 249

FIG. 3. Garden Cove, ar low water. Traverses A and B (Fig. 9) were on rock faces in right middle distance. gastropod Laevilittorina caliginosa, the isopod and varied population which is listed in Table 2. Exosphaeroma gigas, and small numbers of the In contrast there is a marked paucity of spe­ polychaetes, Boccardia polybranchia, Cirratulus cies associated with the encrusting coralline cirratus, Platynereis magalhaensis, and Spirorbis where the kelp is absent. Small groups of Spir­ aggregatus. Small colonies of the bryozoan Ba­ orbis aggregata and a few specimens of the rentsia aggregata are present. echinoderms Pseudopsolus macquariensis and KELP ZONE; In this zone the rock surfaces are Stichaster suteri were noted. dominated by the massive stipes of Durvillea LOWER RED ZONE; Below the Durvillea zone antarctica (Fig. 8), many of the plants having in the truly infralittoral is a community com­ holdfasts 40 cm in diameter. Among ·the Dur­ prised of several species of red algae: Ballia cal­ litricha, Ceramium rubrum, Delessaria sp., lridea villea stipes the rock is covered by a layer of sp., Plocamium coccineum, and Schizoseris sp. encrusting red coralline, alga, the upper limit of The brown alga Desmarestia rossi is common which is usually a little below that of the kelp. and (as mentioned above) Rhodymenia fre­ In the few situations (usually protected from quently extends below the kelp zone. The coral­ wave action) where the kelp is poorly developed line covering the rocks continues below LWS the coralline alga is the dominant organism. except in certain silted areas. The laminae of Durvillea are singularly de­ The fauna of this zone is a large and varied void of life. However, the goose barnacle, Lepas one with some animals dependent on a particu­ australis, was recorded from pieces of kelp found lar type of habitat and others distributed widely. on the beach after gales. In Table 3 the common species are listed with The recesses of the holdfasts suppOrt a dense their representative habitats. 250 PACIFIC SCIENCE, Vol. XVI, July 1962

Compal'iso1Z of Traverses same in each case and only minor variations in As would be expected, the vertical dimension density were noted. Chaetangium fastigiatum of each of the dominant zones is influenced by was less dense at A than at B and absent on the the combined effects of the slope of the rock vertical face. Rhodymenia sp. was observed on surface and the exposure to wave action. the exposed rock face at A but not at either of Comparable measurements were made at par­ the other localities. ticular localities and the zonation picture for Two "flat" reefs were surveyed in detail, sta­ each is shown in Figure 9. tion 1 in Buckles Bay being considered as typi­ cal of this type of rock formation, and a gently Garden Cove, traverse A; slope 80°, on the sloping platform near the fishtrap (Fig. 11) as north shore of the cove, near the entrance a contrast to the nearby vertical face (see above). to the cove and exposed to all weather from Station 1 was reasonably protected by off­ the east or southeast (Figs. 3, 10). shore rocks and a wide belt of Durvillea. The Garden Cove, traverse B; slope 60°, on the true tidal levels given in Figure 11 are those north shore of the cove, partially protected recorded from the tide gauge (which was in­ from surf but covered by the wash from stalled on this reef) during its functional period swell. in 1950. Although spread horizontally and modi­ fied by the intrusion of pools and gutters the Fish Trap, vertical; facing southeast, par­ vertical range of the major zones is comparable tially protected by offshore rocks and kelp with that of the steeper reefs (Fig. 9). (Fig. 5). The more protected and gently sloping (l00) The obvious differences at these three locali­ reef near the fish trap showed a simplified ver­ ties are: sion of the typical zonation pattern with only 1) On traverse B the extension upwards of the dominant organisms present in any density, the zones above MSL, their greater width and and an expanded "bare" zone (Figs. 9, l1),'which greater density. The raising of effective tidal supported more gastropods, Macquariella hamil­ heights at the upper levels of sloping shores has toni, than usual as well as the typical Siphonaria been discussed by Endean, Kenny, and Stephen­ lateralis. son (956). The gullies crossing this reef were awash and 2) The wider "bare" zone and its sparser many small Durvillea plants grew there. flora and fauna on the vertical slope. The Porphyra and Rhizoclonium zone was 3) The deeper extension, below LWS, of the protected from all but the heaviest weather and kelp on traverse A due, presumably, to the normally was dampened by fine spray only. heavier surf in this area. Station 2 (Fig. 9) was a vertical rock face on The species composition was essentially the the shoreward side of a reef near station 1 and

TABLE 2

FAUNA OF Durvillea HOLDFASTS (Arranged alphabetically)

_____V_E_R_y_C_O_M_M_O__N + C_O_MMO:_N : --=P-=-R=E=SE=N:..:-T~, _ Aphroceros sp. Hyale novae-zealandiae Barentsia aggregata Exosphaeroma gigas Laevilittorina caliginosa Hyale hirtipalma Macquariella hamiltoni las pubescens Nemertopsella marri Jassa falcata Spirorbis aggregata Munna maculata Paramoera schellenbergi Procerodes ohlini Psettdopsolus macquariensis Macquarie Island-KENNY and HAYSOM 251

FIG. 4. Buckles Bay with Station 1 reef in the middle distance. sheltered from surf but washed by spent waves. Durvillea antarctica plays an important parr The main variation from the typical pattern was in protecting the reefs from the full force of the the obliteration of the "bare" zone by the over­ surf, the floating fronds damping the breaking lapping of the Porphyra and Red belts. of the swell. However, during storms the lami­ nae of the kelp are whipped against the rock Role of Kelp surfaces and at some localities this appears ro be The large brown algae are the most conspic­ a factor contributing to the sparseness of the uous feature of the Macquarie 1. marine flora. "bare" zone fauna. Extensive beds of Macrocystis pyrifera occur Rock Pools offshore in several fathoms of water, and the island is fringed by Durvillea antarctica attached Many rock pools were examined and in gen­ ro reef edges and offshore rocks (Figs. 8, 12). eral revealed a group of organisms similar [Qo Macrocystis, growing in deeper water, escapes that of the surrounding rocks, with the addition much of the force of wave action and is char­ of species from lower tidal levels. acterised by a more slender and longer form The pools maintained by splash or the high than Durvillea. The only conspicuous organisms seas of srorms present a habitat of extreme con­ on the fronds of Macrocystis were extensive ditions. During the winter months they freeze colonies of the hydroid Orthopyxis platycarpa, on some occasions. At any time of the year the but the tangled holdfasts showed evidence of a decomposition of broken kelp, hurled onto the rich infralitroral fauna. The fauna of Durvillea reefs by storms, must cause considerable varia­ holdfasts has been listed above (Table 2). tionin chemical composition of the pools. Dur- 252 PACIFIC SCIENCE, Vol. XVI, July 1962

ing the summer period they are fouled by ele­ molluscs, especially Siphonaria, could also con­ phant seals, Mirounga leonina. tribute. A pool, 5 m wide and 25 cm deep at approxi­ Predators may play an importaht part in lim­ mately HWS near station 1, was examined on iting the population density of some species. several occasions. At the end of the 1948-49 The introduced, flightless "woodhen," Galli­ summer (l7.ii.49) there was present a rich 1'allus australis, and the Dominican gull, Larus population including Viva, Enteromorpha, dominicanus, forage in the upper algal zones for Rhodymenia, small patches of encrusting red food. Examination of stomach contents of coralline alga, Macquariella hamiltoni, Siphon­ Dominican gulls shows that the gastropod Can­ aria lateralis, Exosphaeroma gigas, Hyale novae­ tharidus coruscans forms a major part of the diet zealandiae, Marionina antipodum, turbellarians, of these birds. The denser populations of these and copepods. molluscs occurred where there was a heavy algal The same pool at the end of the 1949-50 cover, presumably protecting them. summer (9.iii.50) was full of rotting kelp and Stomach contents of the kelp-inhabiting fish sea elephant faeces with no trace of the former N otothenia macrocephala included a wide va­ population. riety of crustacean and polychaete fragments. Biotic Factors DISCUSSION As mentioned above, the movement of the kelp fronds is a probable factor in delineating In contrast to Australian coastlines the Mac­ the "bare" zone, but grazing of this area by quarie I. rocky shore is dominated by algae;

TABLE 3 FAUNA OF THE LOWER RED ZONE·

ABUN- ABUN- ORGANISM HABITAT DANCE ORGANISM HABITAT DANCE Aphroceras sp...... _...._ A,B 4 Munna maculata__ ._____ ..... __ .____ E 2 Myriothela meridiana._ .. ____ . __ B 4 Ta.~ais litoralis.. ____ ...... _..._.. E 2 HaliantheUa kerguelensis...... C 3 Hyale novae-zealandiae...... __ B,C,D,E 2 Parantheopsis cruentata.. ______C 3 Jassa falcata...... ____ .. ____ . B,C,D,E 2 Procerodes ohlini.. ______D,E 3 Paramoera scheUenbergi.. __ .... B,C,D,E 2 NemertopseUa marri...... B,D 4 Halicarcinus planatus.. _____ ..... B 4 Line1U scotti. ___ ...... _...... B,D 4 Pycnogonum platylophum.... D 4 Nematoda (unidentifiedJ ______I C 2 T anystylum neorhetum...... D 4 Arenicola assimilis.. ______.____ ... D 4 T anystylum styligerum...... D 4 Boccardia polybranchia...... C 3 CantharidttJ coruscans. ______A,B 3 Cirratulus cirratus.. ______C 3 Laevilittorina caliginosa.... ____ A,D,E 2 Fabricia alata..... __ ...... C 3 MacquarieUa hamiltoni...... A,D,E 2 Nereis kerguelensis.. ______... C 3 NaceUa delesserti...... __ .____ A,B 3 Platynereis magalhaensis...... C,D 2 Gaimardia trapesina_ ...... E 2 Spirorbis aggregatus...... ____ . A,B 3 Kidderia pusiUa...... __ C 2 Marionin.ti werthi...... C 3 Bal'entsia aggregata...... B 4 Copepoda (unidentifiedJ ______. C,D,E 2 Stichaster suteri...... A,B 4 Cassidinopsis emarginata...... E 2 Pseudopsolus macquariensis.. A,B,C 2 Exosphaeroma gigas.... __ ... _____ D,E 1 Molguia novae-zealandiae..... C 2 las pubescens______.______...... D,E 3

• Letter symbols refer to type of habitat; numerical symbols to relative density of species; organisms are listed alphabetically within taxonomic groups. A, upper surface of rocks 1, very common B, crevices 2. common C, sand and mud 3, present D. under embedded stones 4, occasional specimens E, algal fronds Macquane Island-KENNY and HAYSOM 253

FIG. 5. Fish trap station, showing dense masses of Durvillea.

Porphyra, Durvillea, and various red algae being With the absence of these zones and the climatic the most obvious organisms. The greater role conditions of frequent storms, high winds, and played by the algal elements of the biota in large spray areas, it is difficult to correlate the higher southern latitudes has been noted by Macquarie pattern with the generalised plan of other writers, like Bennett and Pope (1952), either Stephenson and Stephenson (1949) or Guiler (1960), and Knox (1960). The observa­ Womersley and Edmonds (1952). tions in this report suggest that Macquarie ap­ Doty (1957), in his review, comments on the proaches the extreme condition of this latitudi­ intermingling of the terrestrial and truly littoral nal progression in the Australian sector of the flora in areas of heavy spray, and equates the . Other islands in comparable laminarian and Delesseriaceae zone of the North southern latitudes, Kerguelen I. and Heard I., Pacific with the infralittoral zone of Stephenson show a similar algal predominance (personal and Stephenson (1949). A similar situation pre­ observation, R. K.). vails at Macquarie I. Stephenson and Stephenso·n (1949) have sug­ A littorinid and a balanoid zone are re­ gested that a "littorinid-balanoid-Iaminarian" corded from Tasmania (Guiler, 1952) and south­ zonation is of almost universal occurrence on ern New Zealand (Batham, 1960), although in cool temperate exposed rocky shores although the former area the barnacles were considered the species present may vary. The kelp zone at to be of reduced importance, compared with Macquarie can be considered as the "laminaria" lower latitudes. Powell (1955) reports littorinids of the typical plan. Nothing equivalent to the from Auckland I. "littorinid" or "balanoid" zones was observed. Mussels, common on coasts of southern Aus- 254 PACIFIC SCIENCE, Vol. XVI, July 1962 tralia (Bennett and Pope, 1952), New Zealand Zealand (Batham, 1956), and South American (Batham, 1960), and Chile (Guiler, 1959), and shores (Guiler, 1959) is not represented at Mac­ recorded by Powell (1955) at Auckland I., were quarie, although small ascidian (Molgula) com­ not collected in this survey. munities were noted at the lower levels of the The tube worm, Spirorbis aggregatus, occurs shore. in patches and cannot be considered ecologically On the Pacific coast of South America species equivalent to the zones of tube-building poly­ of Siphonaria, Nacella, and Lessonia are impor­ chaetes in warmer latitudes (Dakin, Bennett, and tant organisms (Guiler, 1959) and these genera Pope, 1948; Bennett and Pope, 1952). In New are represented in the collections from Mac­ Zealand the tube-building polychaete, Poma­ quane. toceras coeruleus, is less common in the south, Of southern hemisphere localities at which near Dunedin (Batham, 1956), than near Christ­ comparable investigations have been made, the church (Knox, 1953). west coast of South Africa (Isaac, 1937) shows The Pyura· zone, so prominent on Australian a general format similar to Macquarie 1. There, shores (Dakin, Bennett, and Pope, 1948), New the shore is alga-dominated and shows zones

0 0 Wireless .0 . 0 . 0 • Hill 0 C) 0 Q • 0 () •

Hasselborough Trap Bay •

Buckles Bay North

Main Plateau o 500 1000 .' Yards ... Sites Mentioned in Text

FIG. 6. Isthmus area, Macquarie Island, showing localities mentioned in text. Macquarie Island-KENNY and HAYSOM 255

FIG. 7. Surf conditions at Station 1 during heavy weather.

of Porphyra, Chaetangium, a "bare" area with Arctic. If the Macquarie fauna is examined on limpets, Iridaea, a Champia-Corallinaceae com­ a similar basis, the absence of barnacles and the plex, and an Ecklonia-Laminaria-Macrocystis concentration of molluscs at the lower tidal belt. This coast is affected by the Benguela cur­ levels suggest that this locality is near the zoolo­ rent and, although in latitude 31 0 S, is essen­ gical limit of the Antarctic, with respect to the tially a cold water environment. littoral fauna. In the northern hemisphere there have been This suggestion is borne out by the seasonal many reports of rocky shore zonation in lati­ movements of the Antarctic Convergence from tudes similar to Macquarie I. For example, north to south of the island and the recorded Kitching (1935) noted the typical littorinid­ inshore sea temperatures (see above). Vaughan balanoid-Iaminaria pattern on the coast of Ar­ (1940) delineates the "subpolar" water mass as gyll, in latitude 56 0 N, but this area is not iso­ varying from 5 to 10 C, which is a higher range lated by a sea barrier (Crisp and Southward, than that recorded at Macquarie I. 1953) and apparently is subjected to less rigor­ However, Southward (1958) remarks that the ous climatic conditions. differences between the Antarctic and Subant­ Madsen (1936), discussing the shore fauna of arctic littoral fauna result from varying degrees East Greenland, recorded no barnacles or mol­ of tolerance of cold and ice action rather than luscs, presumably due to the short duration of from particular differences in the species com­ ice-free conditions. He considered that the position of the zonation plan. northern limit of Balanus balanoides and littoral Table 4 lists 58 species of plants and animals molluscs formed the zoological boundary of the collected in this survey for which definite dis- 256 PACIFIC SCIENCE, Vol. XVI, July 1962 tribution records have been traced. Of this num­ Magellan areas may in fact have a wider Sub­ ber, 8 species appear to be endemic to Mac­ antarctic distribution and further collections quarie, 14 have been noted in the New Zealand may increase the listing of Antarctic littoral area, 12 have been recorded from Kerguelen, 18 forms. have wide Subantarctic distribution, and 6 have From these collections it would appear that been recorded from Antarctica. It is possible the littoral biota of Macquarie I. is of a general­ that species listed from the Kerguelen and ised Subantarctic type (Adenocystis, Durvillea,

TABLE 4

BIOGEOGRAPHIC DISTRIBUTION OF SOME RECORDED SPECIES

c c "" ~ ."'" ."'" a U '"0 '"0 u u "" E:: "" ;:: ":Ii u ":Ii u E-< E-< C Ul ." C Ul ." '"E-< Cl '"E-< Cl Z Z "" z z z I "" Z ." '"u ."..., ." ." '"u ." ." '" p E:: '" p ~ E:: ~ ~ u c g u c ..., Ul'" ..., Ul'" ." < ." "" ." <:;0 ." "" :;0 '"E-< ." '"E-< Z ...,." Z '"Z ...,'" u Z '" '"' '" u '"'Z 0 ." ..., Z 0 ." ..., ." E:: ." ;:: "" ..., § "" ..., ~ u § p p ..., e: u p p ..., '" Ul :;§ Ul'" "" P f:1i '" "" ." SPECIES "'" U"'" ." SPECIES Cl .,; u '"E-< Cl .,; '"E-< Z " ""j ~ ""Z " "" ~ z " z ~'" "::s Z ~'" U ." z "" U ." "" "" "'" --"" ------"'" I-- Enteromorpha i1;testir;alis..... X Platynereis magalhaensis . X Adenocystis utricularis...... X Polycirrus kerguelensis . XX X Desmarestia willi...... X Potamilla antarctica . X DurlJillea antarctica...... X Scolelepsis cornifera . X Lessonia variegata...... X Lumbricillus macquariensis.. X Macrocystis pyrifera...... X Marionina antipodum . X Ploccameum coccineum...... X M. werthi . X Halcampoides kerguelensis.. X Exosphaeroma calcareous.. . X Procerodes ohlini...... X E. gigas.. . X Lineus scotti...... X las pubescens . X Barentsia aggregata...... X Jatzira neglecta . X Hemiarthrum hamiltonorum. X Tanais litoralis.. . X H. setulosum...... XX Acontiostoma marionis.. . X Plaxiphora aurea...... X Hyale hirtipalma . X Terenochiton fairchildi...... X H. novae-zealandiae . X Cantharidus coruscans...... X Jassa falcata . X Laevilittorina caliginosa...... X Nannonyx kidderi : . X Macquariella hamiltoni...... X Parajassa tristanensis.. . X Margarella macquariensis...... X Paramoera macquariae X Nacella delesserti...... X P. schellenbergi . X Siphonaria lateralis...... X Pycnogonum platylop;~1Im . X Gaimardia smithi...... X Tanystylum styligerum . X X G. trapesina...... --_ ...... X A!sterina hamiltoni . X Kidderia macquariensis...... X Sporasterias directa . X K. pusilla...... X S. mawsoni . X Tawera mawsoni...... X S. sphoerulata . X Arenicola assimilis...... X X Stichaster suteri . X Cirratulus cirratt/s...... X Pseudopsolus macquariensis.. X Nereis kerguelensis...... X Harpagifer bispinis.. . X Notomastus latericeus...... X XII Notothenia macrocephala . X Macquarie Island-KENNY and HAYSOM 257

FIG. 8. Durvillea zone, at low water.

Exosphaeroma, ]assa, Lessonia, Macrocystis, With one exception, Ploccameum coccineum, Notothenia, Siphonaria, etc.). There is a definite the species reported from Antarctica are also relationship between Macquarie 1. and the Ker­ known from Subantarctic localities other than guelen region, nearly 20% of the species listed Macquarie 1. being present in these two localities. This simi­ larity has been noted previously, by Dell (1952) Species Recorded discussing the fauna in general and by Powell The species collected in this survey are listed (1951) with reference to the molluscs. below. In some groups the list is incomplete The affinities with the New Zealand area due to identification problems. Many species (see also Chilton, 1909) are equally strong; and previously recorded from Macquarie 1. were not Powell (1955) lists Cantharidus coruscans and represented in these collections. Margarella macquariensis, both previously con­ sidered endemic to Macquarie 1., from the Chlorophyceae Auckland Is. Cladophora spp. The endemic species (15 % of those listed) Enteromorpha bulbosa (Suhr) Kuetz are molluscs and echinoderms. Ekman (1953) Enteromorpha intestinalis (L) has commented on the endemic nature of the Monostroma sp. echinoderm fauna, and to a lesser extent the Prasiola sp. molluscan fauna, and uses this distribution to Rhizoclonium sp. suggest that Macquarie 1., occupies a position VIva lactuca (L) quite distinct from the New Zealand Subant­ Phaeophyceae arctic islands. Adenocystis utricularis (Bory) Skottsberg 258 PACIFIC SCIENCE, Vol. XVI, July 1962

Chordaria dictyosiphon (Harvey) Kuetz Rhodymenia spp. Desmarestia willi Riensch Schizoseris sp. Durvillea antarctica (Chamisso) Hariot. Lichens Ectocarpus confervoides (Roth) Le Jolis Verrucaria sp. Geminocarpus geminatus (Hooker and Mosses Harvey) Skottsberg Ceratodon purpureus (Hedwig) Lessonia variegata Agardh MZielleriella crassifolia (Hooker) Macrocystis pyrifera (L) Agardh Angiosperms Scytosiphon lomentaria (Lyngbye) Agardh ColobanthziS muscoides Hooker Rhodophyceae Cotula plumosa Hooker Acrosiphonia pacifica Kiitz Poa foliosa Hooker Ballia callitricha (Agardh) Montagne Puccinellia macquariensis (Cheeseman) Bostrychia vaga Hooker and Harvey Porifera Cerami'(-tm rubrum (Hudson) Agardh Aphroceras sp. Chaetangium fastigium (Bory) Agardh Corallina officinalis (L) Coelenterata Delessaria spp. Hydroidea Hildenbrantia sp. Myriothela meridiana Briggs lridea oborata (Klitz) Setchell and Gardner Orthopyxis platycarpa Bale Melobesia sp. Actiniaria Porphyra columbina Montagne Halcampoides kerguelensis Kwierniewski P. umbilicalis (L) Agardh Parantheopsis cruentata Couth Plocamium coccineum Lyngbye Platyhelminthes Polysiphonia anisogona Hooker an:! Harvey Procerodes ohlini (Bergendal)

GARDEN COVE STAT. 1 STAT. 2 FISH TRAP STAT.

15

A B VERT. HORIZ.

10 i I ~ i r 5 n~ n~ u~ p~ pI p~

0 "... .c Co en ""'Co "0 :5 Q; 2; ~ ~ ....:I ~ ~ ~ ~ ....:I ~ ~ "" ....:I ~ ~ "" ....:I "" ~ "" ....:I "" :3 ~ '" "" FIG. 9. Comparison of zonation at various localities: x-see Fig. 11 solid bar-clense growth open bar-sparse growth Macquarie Island-KENNY and HAYSOM 259

FIG. 10. Garden Cove, at low water. Main traverse area showing Durvillea, "bare" zone, and Porphyra.

Nemertea Kidderia macquariensis Hedley Nemertopsella marri Wheeler K. pusilla Gould Lineus scotti Baylis Tawera mawsoni Hedley Bryozoa Annelida Baremsia aggregata Johnston and Angel Polychaeta Mollusca Arenicola assimilis a/finis Ashworth Amphineura Boccardia polybranchia (Haswell) Hemiarthrum hamiltonorum Iredale and Cirratulus cirratus (Muller) Hull Exogene sp. H. setulosum Dall Fabricia alata Ehlers Plaxiphora aurea Spalowsky Lumbrinereis cingulata (Ehlers) T erenochiton fairchildi Iredale and Hull N ereis kerguelensis McIntosh Gastropoda Cantharidus coruscans (Hedley) N otomastus latericeus Sars Laevilittorina caliginosa Gould Platynereis magalhaensis Kinberg Macquariella hamiltoni (Smith) Polycirrus kerguelensis (McIntosh) Margarella macquariensis Hedley Potamilla antarctica (Kinberg) Nacella delesserti Phillipi Scolelepsis cornifera Ehlers Siphonaria lateralis Gould Spirorbis aggregatus Caullery and Mesnil Lamellibranchiata Syllis sp. Gaimardia smithi Suter T helepus plagiostoma (Schmarda) G. trapesina Lamark Typosyllis sp. 260 PACIFIC SCIENCE, Vol. XVI, July 1962

Oligochaeta Amphipoda Lumbricilltts macquariensis Benham Acontiostoma marionis Stebbing Marionina antipodum Benham Hyale hirtipalma (Dana) M. werthi Michaelsen H. novae-zealandiae (Thomson) Arthropoda ]assa falcata (Montagne) Cirripedia Nannonyx kidderi (Thomson) Lepas australis Darwin Parajassa tristanensis (Stebbing) Isopoda Paramoera macquariae Nicholls Cassidinopsis emarginata (Guerin) P. schellenbergi Nicholls Exosphaeroma calcareous (Dana) E. gigas (Leach) Brachyura las pubescens (Dana) Halicarcinus planatus (Fabricius) ]anira neglecta Chilton Pycnogonida Limnoria antarctica Pfeffer Pycnogonum platylophum Loman Munna maculata Beddard T anystylum neorhetum Marcus Tanais litoralis Vanhoffen T. styligerum Miers

'6 Station 1

4

HW S 2 HW N 0 MSL LWN -2 LWS

5 10 15 20 25 30 Scale in Feet

10 Fish Trap - Horizontal Traverse

'5

{)

RHlZOCLONlUM DURVILLEA

PORPHYRA RHODOMENIA etc. LITHOTH AMNION

FIG. 11. Horizontal zonation. Macquarie Island-KENNY and HAYSOM 261

FIG. 12. Entrance to Garden Cove, viewed from Camp Hill, showing kelp growth.

Acarina Pisces Halozates sp. Harpagifer bispinis (Schneider) Insecta Notothenia macrocephala Gunther Antarctophytosus macquariensis Womersley Aves Antarctopria latigaster Brues Galliralltts australis scotti (Grant) Antarctotachinus crozetensis Enderlein Larus dominicanus Lichtenstein Coelopa macquariensis W omersley Mammalia Ephydra macquariensis Womersley Mirounga leonina (L) Pro canace macquariensis Womersley Echinodermata ACKNOWLEDGMENTS Asteroidea The authors wish to thank Mr. P. G. Law, Asterina hamiltoni Koehler Director, Antarctic Division, Department of Ex­ Sporasterias directa (Koehler) ternal Affairs, Australia, for permission to pub­ S. mawsoni (Koehler) lish this work, and to acknowledge the assistance S. sphoerulata (Koehler) given, in the field, by many members of the Stichaster suteri (Loriol) Division. In particular, thanks are due to Mr. R .. Holothuroidea Dovers for assistance in surveying reef profiles. Pseudopsolus macquariensis (Dendy) Thanks are due to the many people who as­ Chordata sisted with the identification of collections: Mrs. Urochordata I. Christianson (formerly Botany Department, Molgula novaezelandiae (Michaelsen) University of Melbourne), Algae; Dr. W. Daw- 262 PACIFIC SCIENCE, Vol. XVI, July 1962 bin (Zoology Department, University of Syd­ CHILTON, C. 1909. The Sub Antarctic Islands ney), Holothuroidea; Dr. H. B. Fell (Victoria of New Zealand. Canterbury, N. Z. University College, N. Z.), Asteroidea; Dr. O. CRISP, D. J., and A. J. SOUTHWARD. 1953. Iso­ Hartman (Allan Hancock Foundation, Califor­ lation of intertidal animals by sea barriers. nia), Annelida; Miss J. H. Macpherson (Na­ Nature 172: 208. No. 4370. tional Museum, Melbourne), Mollusca; Miss J. Nurse (Canterbury University College, N. Z.), DAKIN, W. J., I. BENNETT, and E. C. POPE. Turbellaria; Dr. S. Prudhoe (British Museum, 1948. A study of certain aspects of the ecol­ Natural History), Nemertea; Dr. K. Sheard ogy of the intertidal zone of the New South (CSIRO Division of Fisheries and Oceanogra­ Wales coast. Aust. Jour. Sci. Res. ser. B, 1: phy, W. Australia), Crustacea. 176. Professor W. Stephenson (Zoology Depart­ DEACON, G. E. R. 1937. The hydrology of the ment, University of Queensland, Australia) Southern Ocean. Discovery Reps. 15: 1. kindly read and criticised the manuscript. DELL, R. K. 1952. In: The Antarctic Today, Wellington, N. Z. SUMMARY DoTY, M. S. 1957. Rocky intertidal surfaces. The major features of the rocky shore zona­ Geol. Soc. Am., Mem. 67: 535. tion pattern at Macquarie I., are described as ENDEAN, R., R. KENNY, and W. STEPHENSON. comprising a lichen zone; a Porphyra zone; a 1956. The ecology and distribution of inter­ "bare" zone dominated by Siphonaria; an upper tidal organisms on the rocky shores of the red algal zone, predominantly Rhodymenia; a Queensland mainland. Aust. Jour. Mar. Durvillea zone; and a lower red algal zone of Freshw. Res. 7: 88. Dellessaria, Iridea, and Desmarestia. The secondary organisms associated with the EKMAN, S. 1953. Zoogeography of the Sea. above zones are listed. London. The zonation pattern is compared with simi­ GUILER, E. R. 1952. The nature of intertidal lar ecological situations in southern Australia, zonation in Tasmania. Pap. Proc. Roy. Soc. New Zealand, South Africa, and South America. Tas. 86: 31. The algal domination of the shore shows much --- 1959. The intertidal ecology of the in common with the west coast of South Africa. Montemar area, Chile. Pap. Proc. Roy. Soc. The absence of barnacles and littorinids sug­ Tas. 93: 165. gests that the Macquarie shore is more typically --- 1960. The intertidal zone forming spe­ Antarctic than Subantarctic. The geographic cies on rocky shores of the East Australian distribution of species shows the island to have Coast. J. Ecology 48: 1. a closer biogeographic relationship to other Subantarctic islands than to the New Zealand ISAAC, W. E. 1937. Studies of South African area. seaweed vegetation, I. West coast from Lam­ berts Bay to the Cape of Good Hope. Trans. REFERENCES Roy. Soc. S. Afr. 25: 115. BATHAM, E. J. 1958. Ecology of a southern KITCHING, J. A. 1935. An introduction to the New -Zealand exposed rocky shore at Little ecology of intertidal rock surfaces on the Papanui, Otago Peninsula. Trans. Roy. Soc. coast of Argyll. Trans. Roy. Soc. Edinb. 58: N. Z. 85: 647. 351. BENNETT, I., and E. C. POPE. 1953. Intertidal KNOX, G. A. 1953. The intertidal ecology of zonation of the exposed rocky shores of Vic­ Taylor's mistake, Banks Peninsula. Trans. toria, together with a rearrangement of the Roy. Soc. N. Z. 81: 189. biogeographical provinces of temperate Aus­ --- 1960. In: Biology of the Southern Cold tralian shores. Aust. Jour. Mar. Freshw. Res. Temperate Zone, ed. Holdgate, Nature 185: 4: 105. 203. No. 4708. Macquarie Island-KENNY and HAYSOM 263

MACKINTOSH, N. A. 1946. The Antarctic Con­ SOUTHWARD, A. J. 1958. Zonation of plants vergence and the distribution of surface tem­ and animals on rocky sea shores. BioI. Rev. perature in Antarctic waters. Discovery Reps. 33: 137. 23: 177. STEPHENSON, T. A., and A. 1949. Universal MADSEN, H. 1936. Investigations on the shore features of zonation between tide-marks on fauna of East Greenland with a survey of the rocky shores. J. Ecology 37: 289. shores of other Arctic regions. Medd. om TAYLOR, B. W. 1955. The flora, vegetation, Gr¢nland 100(8): 1. and soils of Macquarie Island. Aust. Nat. Antarct. Res. Exp. Reps. Ser. B, 2: 1. MAWSON, SIR D. 1943. Macquarie Island, its geography and geology. Aust. Antarctic Exp. VAUGHAN, T. W. 1940. Ecology of modern Reps. ser. A, 5: 1. marine organisms with reference to paleo­ geography. Bull. Geol. Soc. America 51: 433. POWELL, A. W. B. 1951. Antarctic and Sub­ WOMERSLEY, H. B. S., and S. J. EDMONDS. antarctic Mollusca. Discovery Reps. 26: 47. 1952. Marine coastal zonation in southern --- 1955. Mollusca of the southern islands Australia in relation to a general scheme of of New Zealand. Cape Exp. Series, Bull. 15. classification. J. Ecology 40: 84.