Oceanography and Marine Zoology of the New Zealand Subantarctic

44 ECOLOGY OF SUBANTARCTIC ISLANDS

The highest temperatures recorded at Auck- REFERENCES land Is. and at Campbell 1. are much the same, approximately 65°F., and are 12°F. DE LISLE, J. F., 1964. Weather and climate of Campbell higher than that at Macquarie 1. (Table 12). Island. Pacific Ins. Monograph 7: 34-44. The extreme minimum at Macquarie I. is 10°F. lower than that at Port Ross. FABRICIUS, A. F., 1957. Climate of the Subantarctic Islands. In Meteorology of the Antarctic, ed. Van Sea temperatures at Campbell 1. are about Rooy. Weather Bureau, South Africa. 3°F. warmer than the earth temperatures at 12 inches in mid winter and about I!OF. colder FALLA, R. A., 1948. The outlying islands of New Zea- in midsummer (Table 13). land. N.Z. Geographer 4; 127-154. HITCHINGS, M. G., 1949. Campbell Island. A subantarc- TABLE 13. Mean monthly earth and sea tic meteorological station. Weather 4: 389-92. temperatures. Earth temp. Sea temp. MARSHALL, 0., 1909. The Meteorology of Campbell at 1 foot Campbell Auckland Island. In The Subantarctic Islands of New Zea- Campbell L I. Is. land, ed. C. Chilton. Philosophical Institute of Jan. 51.0 49.5 50.8 Canterbury. 2 vols. Feb. 50.3 49.t 51.0 Mar. 48.5 48.5 50.5 MAWSON, D., 1915. The home of the blizzard. Heine- man, London. 2 vols. A~ ~ ~ ~ May 43.2 45.3 47.4 NEWMAN, W., 1929. Tabulated and reduced records of June 40.3 43.2 46.1 the Macquarie Island Station. In Aust. Antarc. July 39.5 42.6 45.7 Exp. 1911-14 Sci. Rep., Ser.B., 3, Sydney. Aug. 40.0 42.9 45.1 Sept. 41.3 43.7 46.0 N.Z. METEOROLOGICAL SERVICE, 1963. Summaries of Oct. 43.4 44.6 46.8 radiosonde data 1956-61. N.Z. Met. S. Misc. Pub. Nov. 46.5 46.1 47.9 119. Dec. 49.6 48.5 49.2 NOTE. Sea temperatures for Campbell 1. taken monthly ROBERTSON,N. G., 1964. The frequency of high intensity in the enclosed waters of Tucker Cove, 1943-55, rainfalls in New Zealand. N.Z. Met. S. Misc. Pub. and for Auckland Is. at Port Ross, 1941-45. tt8.

OCEANOGRAPHY AND MARINE ZOOLOGY OF THE NEW ZEALAND SUBANTARCTIC

ELLIOT W. DAWSON

N.Z. Oceanographic Institute, D.S.l.R., Wellington-

The Subantarctic Islands of New Zealand, regions washed by 'subantarctic water', in comprising Bounty and Antipodes, Auckland the hydrologist's use of the term. Undoubtedly and Campbell, Macquarie and the Snares, lie these more northern regions show the effects within the Subantarctic zone of surface waters, of subantarctic waters and so should be dis- bounded on the north by the Subtropical Con- cussed with the New Zealand Subantarctic. vergence Region and on the south by the Similarly, Macquarie 1., lying close to the Antarctic Convergence about the latitude of Antarctic Convergence, should be included Macquarie 1. (Fig. 1). because it shows the mixed influence of Antarctic and Subantarctic waters. Whether the Chatham Is., the Snares, and even the southern New Zealand mainland . The Society is grateful to the N.Z. Oceanographic should be considered 'subantarctic' may be Institute for a contribution towards the cost of pub- resolved by defining as 'subantarctic' any lishing this paper.-ED. DA\VSON: OCEANOGHAPHY AND MARINE ZOOLOGY 45

- FIGURE 1. The Subantarctic Region and associated physical features of the Southern Ocean.

HISTORY OF MARINE EXPLORATION were made by naturalists as passengers in the N.Z. Government vessels Stella, Tutanekai and Oceanographic and marine biological ex- Hinemoa, under Captains Fairchild and ploration of the Subantarctic region began Bollons, both of whom were amateur naturalists about 1840 with the great voyages of D'Urville, of some experience. The real foundation of Ross and Wilkes and continued through tbe our knowledge of the marine fauna of the work of the French Transit-of-Venus Expedi- New Zealand Subantarctic was laid by the tion to Campbell Island in 1874. As well as 1907 expedition and was later added to by the the famous 1907 Expedition of the Philoso- results of Dr. Th. Mortensen's cruise in the phical Institute of Canterbury, irregular visits Amokura as part of his Pacific Expedition +6 ECOLOGY OF SUBANTARCTIC ISLANDS

(191+-1916), and by the war-time Cape rounding surface water moves and at which Expedition, the results of which appear in the it sinks. This movement is related to differ- D.S.1.R. Cape Expedition Bulletin and latterly ences of temperature and salinity. North of in the Records of the Dominion Museum. Dell the Antarctic Convergence, which lies between (1952) and Knox (1960) have both discussed 52°S. and 62°S., the Subantarctic Region the zoogeographical affinites and relationships extends to meet the warmer Subtropical Water of the marine fauna of the New Zealand in the 'Subtropical Convergence Region' Subantarctic based on reports of these earlier between 35°5. and +7°5., which extends expeditions which were, however, largely further north in summer than in winter and confined to the immediate surroundings of the varies from a broad zone near the west of islands. The nature of the fauna, sediments New Zealand to a narrower and better defined and bottom features of the ocean around and zone to the east. There are two types of Sub- between these islands were until recently still antarctic Water with differing salinity: unknown. Australasian Subantarctic Water mainly south of Australia and New Zealand, and Circum- N .Z. OCEANOGRAPHIC INSTITUTE IN THE polar Subantarctic Water further south. SUBANTARCTIC The New Zealand Oceanographic Institute An interesting feature of subantarctic hydro- has in recent years extensively sampled and logy is the intrusion from the Tasman Sea west explored the sea floor surrounding the Sub- of New Zealand of water warmer and more antarctic islands, across the Campbell Plateau, saline than the Subantarctic Water to the east and along the Macquarie Ridge from the New and south. This intruding water is cut off from Zealand Shelf to the Balleny Is. (Dawson 1963, the Subtropical Water to the north by the 196+). upwelling of cold and weakly saline water off Otago. This, and the existence of the bound- The first cruises were largely concerned ary between Circumpolar and Australasian with physical oceanography. Later cruises in Subantarctic Waters, with its associated front H.M.N.z.S. Endeavour (1959, 1963, 1964, along the southern edge of the Campbell 1965), H.M.N.Z.S. Rotoiti (1961, 1962) and Plateau, are important in the marine ecology m.v. Taranui (1962) have concentrated on of the Subantarctic. benthic work. These stations, almost 300 in all, from deep water to intertidal shore collec- Garner (1959), Burling (1961) and Hout- tions, and a further 100 stations on the Chatham man (in press) have discussed the detailed Rise and 400 on the New Zealand Shelf, physical oceanography but ecologists are con- provided a substantial basis on which to begin cerned only with those factors influencing an ecological analysis of the marine benthic animal or plant life. There are still gaps in fauna of the New Zealand Region in general our knowledge of Subantarctic hydrology; for and of the Subantarctic in particular. example, about bottom temperatures and the The sediments of the Subantarctic region detailed effects of submarine topography on are being ana lysed and, apart from dark water movements. Since there are even more volcanic sands and pebbles off Macquarie and gaps in our knowledge of the life-history and Antipodes Is. and the rocky pebble-strewn high behaviour of most marine animals, we may not spots of the Macquarie Ridge, the general pat- yet know what questions to ask the physical tern is of a bryozoan shell sand, with areas of oceanographer nor how he should frame his plain sands on the shelf of each island out to answers to help interpret ecological phenomena. about 100 fathoms; this gives way to the east on the Campbell Plateau to a widespread mass GEOGRAPHY AND GEOLOGY of Globigerina ooze at an average depth of 300 Campbell 1. and the Auckland Is., the fathoms. Both sediments support a rich animal shallow Pukaki Bank, and a recently recog- life, the ooze being of particular interest nised, unnamed submarine volcanic cone, are because of its populations of archibentha] all high spots on an extensive submarine flat, molluscs and echinoderms. the Campbell Plateau, with an average depth of about 300 fathoms (Fig. 2). The Bounty OCEANOGRAPHY and Antipodes Is. to the east form a platform A 'convergence' may be defined as a point of their own separated from the Campbell or line on the sea surface towards whkh SUy- Plateau by depths of about 700 fathoms SOUTH FIJI BASIN

TASMAN BASIN

~ BOUNTY TROUGH

The New Zealand Shelf (a-200m)

The New Zealand Slope or Archibenthal (200- c.IOOOm) Contours in metres

FIGURE 2. The bathymetry of the New Zealand Region, including the New Zealand Sub- antarctic. 48 ECOLOGY OP SUBANTARCTIC ISLANDS

and from each other by more than 500 Forsterian marine province which meets the falhoms. Macquarie 1. is'linked to the New Antipodean near Stewart 1. Obviously such Zealand land mass by a long intermittent sub- provinces cannot be :delimiled by arbilrary marine ridge (Brodie & Dawson, in press) lines and, so long as they are believed to be which extends to the south towards the Balleny correlated with hydrological zones, their Is. on the fringe of the Antarctic Continent. boundaries must be thought of as zones them- Macquarie I. rises above the sea as an selves, varying with time. elongated part of this ridge. There are problems at this stage of ecological The geological history of these islands is analysis in identification of many of the impor- essentially a story of volcanism, erosion, glacia- tant marine groups and there is little com- tion, submergence and deposition. Geological parative material or information from critical accounts of Macquarie 1., of Campbell 1. and areas in the non-New Zealand Subantarctic. of the Snares are available by Mawson (1943), Nevertheless, certain broad features of the Oliver et al. (1950) and Fleminl'( et al. (1953). marine zoology of the New Zealand Sub- The geology of the Auckland Is. has not yet antarctic have emerged and the following been described in detail; but an account of the principal faunal elements can be recognised: geology of the Bounty-Antipodes region is 1. Widespread circum-Subantarctic genera. being prepared (D. J. Cullen, pers. comm.). The over-all geological history forms an 2. New Zealand species, widespread around interesting background to the formation of the the mainland, ranging into the Sub- sediments and submarine features on which antarctic. marine benthic animals live. 3. Endemic subantarctic species of New Zealand genera. BIOGEOGRAPHY 4. Local representatives (subspecies) of polytypic New Zealand genera, parti- The New Zealand Subantarctic Region as a cularly amongst the Mollusca. whole has been regarded as a separate marine faunal province (see Finlay 1925, elaborated Animals within these categories have been by Powell 1951, 1961 and Dell 1962). This specified by Dell (1952) and by Knox (1960, region, which excluded Macquari¤ I., was 1963), but a few other examples, from the originally named the Rossian Province but Echinodermata, will be discussed later to later changed (Powell 1951) to Antipodean, illustrate particular points of interest in sub- preventing confusion with the Rossian subantarctic ecology. province of the Antarctic (cf. Knox 1960, 612). Up to the present, the characteristics of the MACQUARIE ISLAND marine fauna of the Antipodean Province have Macquarie I. was separated from the Anti~ been based on what is largely Cape Expedition podean Province by Powell (1955, 1957), material collected on the islands themselves, largely influenced by the molluscs of a single supplemented by the reports from the Morten- haul (Station 83) of the B.A.N.z.A.R. Expedi- sen Pacific Expedition. Cape Expedition reports tion in 38 fathoms off Lusitania Bay. He include Fell (1953) on the Echinodermata, considered that Macquarie lay within the Powell (1955) on the Mollusca, and Parrott Kerguelenian Province which also included (1958) on fish. Mortensen's own reports on Kerguelen I., Marion, Prince Edward, and the Echinodermata (1921, 1925) are also the Crozet Islands. The faunal affinities of important. the Kerguelenian Province are said to be with The biogeographical relationships of various the Antarctic although there is a high endemic species collected by these earlier expeditions element. have already been summarised by Dell (1952) Dell (1964) has recently shown that the and by Knox (1960, 1963) in reviews which 42 species of marine Mollusca so far recorded illustrate the status quo of subantarctic bio- from Macquarie I. represent a faunal mix- geography until the recent work discussed here. ture derived from various sources, and he has Reports on the molluscs of the Snares (Dell suggested that Macquarie's inclusion within 1963a) and on the benthic fauna of Foveaux the Kerguelenian Province loses sight of the Strait (Dawson, in press) help to define the very distinctive character of the Macquarie D,HVSON: OCEANOGRAPHY AND MARINE ZOOLOGY 49 fauna. Dell's analysis shows the following ". . . the Snares seem to be related more closely relationships: Kerguelenian, 12%; circum- to the New Zealand mainland than to the other subantarctic islands; for, of the four genera known Subantarctic, 12%; Heard Is., 5%; New Zea- to have species there. . . two are represented on land, 5%; of no particular affinity, 2%. The the New Zealand mainland by identical species, recent collections of Echinodermata made by and two others occur in the subantarctic islands the N.Z. Oceanographic Institute off Macquarie as well as the New Zealand mainland, identical species being involved in each case. There are no show a much stronger link with New Zealand. Snares echinoderms which are shared with the other subantarctic islands and not also shared with By contrast, in a report on the intertidal the New Zealand mainland. Hence. . . the rocky shore fauna of Macquarie 1., Kenny and northern boundary of the Rossian marine province Haysom (1962) conclude that Macquarie is would fall south of the Snares; the latter would form part of the Forsterian, with south Otago and biogecgraphically closer to the Antarctic Stewart Island." Province than to the New Zealand Subantarctic region. Their analysis of 58 species is: 8 Since 1953, a further six species of asteroids, endemic, 14 New Zealand, 12 Kerguelen, 18 four of ophiuroids, nine of echinoids, and two wide-ranging Subantarctic, and 6 Antarctic. of holothurians have been taken by the N.Z. This analysis warrants a little nlOre caution Oceanographic Institute from the Snares, and than Kenny and Haysom's generalisation and from shore collections. may further support Dell's recognition of the With the exception of the holothurian distinctiveness of the Macquarie fauna. Stereoderma leoninoides, all are shared with A more detailed analysis of the marine fauna the New Zealand mainland as well as with of Macquarie is difficult because the available the subantarctic islands. The close resemblance collections have been taken from different between the intertidal molluscs of the Snares localities or different environmental niches, so and those of Stewart I. is further evidence of that few of the same species have been taken the link between the two faunas (Dell 1963a). by successive collectors. This is especially so The Snares undoubtedly lie in an interesting among the echinoderms~ for example the marginal region. species of the starfish genus H enricia (Fig. 5). "There seems little doubt that there is a well defined faunal break between the southern islands The conclusion, that Macquarie 1. has a of New Zealand (Campbell, the Aucklands, Boun- mixed fauna, is certainly what would be ties and Antipodes) and the New Zealand mainland including the Snares. At the same time many expected from its hydrological position and of the really characteristic Antipodean genera and is supported by other evidence from the species cross this faunal break and extend to a Echinodermata. varying degree to the north" (Dell 1962; 50).

THE SNARES MARINE FAUNA The northern limit of the New Zealand So far, the systematics of only a few groups Subantarctic (or Antipodean Province) lies, of marine bottom-living animals of the New according to the hydrological definition of Zealand Subantarctic are sufficiently well Subantarctic, in the region of the Snares Is. known to make possible the compilation of To exclude these islands fronI the Subantarctic faunal lists and the plotting of species distribu- is no more reasonable than to exclude tions. Two such groups are the Mollusca and Macquarie I. at the other extension of the the Echinodermata. New Zealand Subantarctic Region. MOLLUSCA Subantarctic Water certainly influences the Snares Is. and their surrounding shelf. A sub- The Mollusca of the shallower parts of the antarctic element is present in their fauna, New Zealand Subantarctic, in particular of the which may be traced to a varying extent in shelf around each group of islands, are cur- Stewart 1. and the southern New Zealand rently being examined and some additional mainland. information to that given by Powell (1955) is already available. For instance, the Four species of echinodenns taken on the gastropod Cymatona kampyla (Watson 1885), shores of the Snares led Fell (1953: 108) to originally described from "Challenger" station conclude: 164B off the New South Wales coast and since 50 ECOLOGY OF SUBANTARCTIC ISLANDS recorded by Dell (1956) from the Chatham TABLE 1. Number of species and biogeographic Rise, off the east Otago coast, and in 200-300 distribution of malor groups of echinoderms fathoms in Palliser Bay, is now known to be (excluding Ophiuroids and Crinoids) from New common throughout the New Zealand Sub- Zealand subantarctic islands (shelf and shore) antarctic particularly in shallower depths off ann the (;amJ1hell Plateau (archibenthal) the Bounty and Antipodes Is. A very closely O.eu '0 " ~~ allied species, Cymatona tomlini Powell, 1955, ~ ~N,O) u _.e°" 00 " u .e °i"O 0.. .e 0" ""~ occurs off Macquarie 1. The echinoderms and .u E "0 E~ ...... D-"~ " 00 ~ E.... molluscs of the Globigerina ooze of the Camp- o~_E.d~ "- " . " ~.....::: Z'" u.... a> "0" bell Plateau vividly demonstrate the uniformity ~.d -".e "'OJ"~ "0 and extensive range of the archibenthal fauna AUCKLAND IS. " of the New Zealand slopes. Asteroids ..... 11 7 2 2 Echinoids "... 4 4 - - Archibenthal molluscs so far identified from Holothurians 8 6 2 - the Campbell Plateau include Penion benthicola CAMPBELL I. Dell, 1956; Ellicea recens Dell, 1951; Galeodea Asteroids 5 4 1 - triganceae Dell, 1953; Cymatona kampyla Echinoids ".. 2 1 1 - Watson, 1885; Coluzea mariae Powell, 1952; Holothurians ."... 4 3 1 - Coluzea altocanalis Dell, 1956; Otukaia blacki ANTIPODES IS. Dell, 1956 (formerly Alertalex); Baryspira Asteroids .--. 2 2 - - bathami DeB, 1956; Pachymelon (Palomelon) Echinoids 4 3 1 - smithi Powell, 1950; Teremelon knoxi Dell, Holothurians 2 2 - - 1956; and Iredalina mirabilis Finlay, 1926; BOUNTY IS. all these are members of the deepwater Asteroids t t - - molluscan fauna recognised by Dell (1956) Echinoids ..... 5 4 1 - Holothurians ---.. 2 1 t - from the Chatham Rise and the New Zealand slopes. These discoveries extend to 52°S., the SNARES IS. Asteroids .." 10 10 - - southernmost limits of this widespread archi- Echinoids 9 9 - - benthal fauna which is also becoming known Holothurians ...... 2 t t - progressively further north, as far as the Bay MACQUARIE I. of Plenty at 35°S. (Dell 1963b). Asteroids 14 6 6 2 Echinoids 1 1 - - One of the biggest question marks lies over Holothurians 6 4 1 the Bounty Trough, the East Otago Slope, and 1 the South Chatham Slope which separate the CAMPBELL PLAT. (Archibcnthal) Campbell Plateau from the Chatham Rise. The Asteroids 17 13 14 , nature of the bottom animals and sediments Echinoids 5 5 - - of these three areas and their possible relation- ,Holothurians Not yet identified - - ship to those of the Campbell Plateau and TOTAL SPECIES 70 48 19 5 Chatham Rise are unknown. NOTE. Asteroids at Macquarie I. include all identifica~ tions of Henricia spp. ECHINODERMS Echinoderms are especially suitable for Longitudinal boundaries are placed at 1580 ecological and biogeographical studies. They and 180°E. The distribution of the echino- are relatively sedentary and usually have a derms (excluding Crinoidea) so far identified limited bathymetrical range, often showing from this arbitrarily defined area is summarised definite sediment preferences and correlations in Table 1. with salinity and temperature. The shortness Several new records have been added to the or absence of a pelagic larval stage is also known species from the Auckland Is. The important in the distribution of certain species. occurrence of the echinoid Apatopygus recens The northern limit of the New Zealand has been confirmed. There also occur on the Sub3ntarctic is here taken at the latitude of the edge of the shelf large numbers of Araeosoma Snares Islands, 48°S., and the southern limit thetidis. This species, previously considered at 58°S., the furthest south from which samples subtropical, is called by Fell (1962a: 84) a are at present available. "dangerous sea urchin" and has been known to DAWSON: OCEANOGRAPHY AND MARINE ZOOLOGY 51 cause painful ,,'.'Ounds in Australia and New viviparous holothurian which ranges from Caledonia. Henricia ralphae and a new species intertidal depths to about 100 fathoms, could of holothurian of the genus Stolus (cr. Pawson presumably only have come to Macquarie in in press) are now recorded. Campbell 1. has such a way. Transport on floating algae has about half the number of species known from been proposed to account for such distributions the Aucklands and all are shared with these (Mortensen 1925); but the problem of moving islands. The holothurian Trochodota dune- against the prevailing water movement is still dinensis is a new record for Campbell 1. real and various species which might havc made use of pelagic drifting of their larvae or Eight species of echinoderms are now known of algal transport arc absent from similar from the Bounty Is. and all except Pseudechinus situations in subantarctic South America and novaezealandiae are new records; they include from other Southern Ocean islands. Other the same new species of Stolus and new species bottom-living holothurians, such as Pso/us of the echinoid Brisaster (d. Pawson in press). antarcticus, found in Magellanic and Antarctic Of nine species from the Antipodes Is. eight South America and Macquarie 1., have pre- are new records. Only three species have been sumably spread along the deep ocean floor or found at both Antipodes and Bounty Is.: along deeper ridges. Apatopygus recens, Pseudechinus novaezea- The echinoids and holothurians of Mac- landiae and Spatangus thor previously known quarie I. are very distinctly related to the only from Foveaux Strait. Of particular interest fauna of New Zealand and the rest of the is a species of Austrocidaris, a genus previously New Zealand Subantarctic, rather than having only known from two species in the Magellanic- marked Kerguelenian or Antarctic affinities. Falklands region. The asteroids are more varied in their relation- Macquarie I. has a rich starfish fauna ships if the presence of a number of species (Clark 1962). Knowledge of the holothurian of Henricia is real and if the occurrence of fauna of Macquarie has been substantially what seems to be a species of Pseudarchaster increased (Pawson in press), and there are from the New Zealand shelf can be confirmed. two new records and three occurrences of The circum-Subantarctic genera Anasterias, species also known from New Zealand. Hymenasler, Smilasterias and Cycethra, and the Antarctic Porania antarctica give a more The only echinoid so far recorded from southerly flavour to the Macquarie fauna, as Macquarie, Pseudechinus novaezealandiae, has might be expected from the situation of Mac- a pelagic larval stage and occurs at all islands quarie on a southwards-extending submarine in the New Zealand Subantarctic (Fig. 3). ridge which straddles the Antarctic Conver- Hydrological circumstances do not favour the gence and is exposed to surface drift from the transport of larvae to Macquarie I. by south- Kerguelenian Province. wards surface water movements. The influence In summary, 68% of the known echino. of the West Wind Drift, which crosses the derms from the New Zealand Subantarctic southern part of the New Zealand Region in a are shared with the New Zealand Shelf or west to east direction, would tend to carry Slope. Endemic species are 27%, and 7% of pelagic larvae well away to the east of New the subantarctic species arc also known from Zealand. Variations in the East Australian the Magellanic or other extra-limital area. Current which impinges on the West Wind Despite Fell's (1962b) otherwise convincing Drift would probably affect such larval drifting hypothesis of echinoderm dispersal through the in the same way. Whereas previously it has Southern Ocean by epiplanktonic drift, the been necessary to postulate a drift mechanism, wind-driven surface layers of the Southern for dispersal to the Macquarie region, a possible Ocean appear to have contributed no species shallow middle depth migration route has now of echinoderms to the New Zealand Suh- been shown to exist. antarctic. However, the seaweed.inhabiting With its known bathymetric range of at holothurians of the genera Traehythyone, least 10 to 150 fathoms, Pseudechinus novae- Stereoderma and Genus may have been dis- zealandiae is much more likely to have reached tributed by this West Wind Drift. Macquarie 1. from New Zealand along the The widespread Australian, Indo-West submarine ridges and sea mounts of the Mac- Pacific and cosmopolitan genera Trochodota, quarie Ridge. Trochodota dUlledinensis, a Chiridota, Spa/angus, Paramaretia, Brisaster SOUTH FI) I BASIN

- ~vv TASMAN BASIN ICHATHAM RISE

-BOUNTY TROUGH

Pukaki Bank

KEY

Pseudechinus a/bocinctus Pseudechinus (Iemingi Pseudechlnus grossu/aria. Pseudechlnus hutton; Pseudechinus novae. zea/andiae Pseudechinus variegatus

FIGUHE 3. The distribution of the echinoid genus P~eudechinu~ within the TVell.' Zealand Region. DAWSON: OCEANOGRAPHY AND MARINE ZOOLOGY 53

and Psolus contribute at least eight species to Goniocidaris umbraculum (Hutton 1872) the fauna; the bathymetrical tolerances of these mainly occurs between 50-200 fathoms in sand species have probably allowed them to migrate and gravelly sand throughout the eastern and along submarine ridges and other bottom southern shelf from Cook Strait to south of features without it being necessary to invoke Stewart 1., although known from 300 fathoms pelagic drift of larvae or epiplanktonic adults in Cook Strait. Recent work on the Chatham as a mechanism. Rise has, however, yielded G. umbraculum from five stations from 240-610 fathoms. These PSEUDECHINUS apparently anomalous records will be discussed elsewhere (Dawson, in progress). G. umbra- The echinoid genus Pseudechinus, exempli- culum has now been found on the Macquarie fied by the widespread species P. novaezea- Ridge and on the Auckland Is. Shelf (Fig. 4). landiae, is worth close ecological examination; It is a brood-protecting species (Mortensen its distribution and sediment preferences on 1926) and its distribution along the Macquarie the New Zealand Shelf have been studied Ridge is comparable with that of Pseudechinus (McKnight, in press). Six species occur in the novaezealandiae, which has pelagic larvae. New Zealand Region, although only P. novaezealandiae is known from the New Zealand Goniocidaris parasol Fell 1958, described Subantarctic. The geographic and bathymetric from 130 fathoms east of the Chatham Is., ranges of all the species are shown in Fig. 3. replaces G. umbraculum in the southern archibenthal of the Campbell Plateau and Pseudechinus novaezealandiae (Mortensen extensively (Fig. 4) on the Chatham Rise. 1921) ranges widely in coarse and fine sands from the southern Cook Strait shelf on the east coast and from off Fiordland on the west ASTEROIDS through the Subantarctic to Macquarie 1., but The starfish of the New Zealand Subantarctic has a limited depth range of 0-168 fathoms have not been fully analysed but an analysis and is found mainly between 12 and 85 of distribution patterns and sediment correla- fathoms. tions of archibenthal species over the Chatham Rise is being prepared (Dawson, in progress) Fell (1962b) presents the hypothesis that and a similar study of the New Zealand Shelf echinoderms are dispersed through the southern species has been made (D. G. McKnight, in circumpolar seas by epiplanktonic drift. He progress) . says of Pseudechinus: ". . . one suspects that the genus set out from New Zealand long ago, A surprising feature of the southern distri- circumnavigated the globe and, like Magellan's butions is the presence of deep-living warm sailors, arrived back whence it started, but water species, such as Benthopecten pentacan- from the other direction." thus Fell 1958 and Zoroaster spinulosus Fisher 1906, known from the Bay of Plenty and the Of the four other species of Pseudechinus Indo-Pacific, and of deep-water starfish of the known to me, two occur in Australia, one off Chatham Rise, Plutonaster knoxi Fell 1958, Marion, Gough, Crozet and Prince Edward Is. Hippasteria tro;ana Fell 1958 and Crossaster and one in southern South America, Falklands, ;aponicus (Fisher 1911). A species of Tristan da Cunha and New Amsterdam. It Pseudarchaster from 200 fathoms off Mac- may be, however, that Pseudechinus novae- quarie 1. is hardly distinguishable from P. zealandiae has not colonized New Zealand from abernethyi Fell 1958, a Cook Strait Shelf the southwest as Fell suggests, but is spreading speCIes. through the southwest from New Zealand to Macquarie along shallow submarine ridge tops. Two unexpected new generic records of starfish from the Campbell Plateau of considerable GONIOCIDARIS biogeographical interest are Lithosoma and Ceramaster, genera previously unknown from The cidarid Goniocidaris is represented by Australasia. The Lithosoma species is probably tbree species in New Zealand (Fig. 4). The new, but related to L. penichra Fisher. new species Goniocidaris magi Pawson 1964, is known from gravelly sand in 30-50 fathoms Lithosoma comprises five known species, all off the Three Kings and may be common in Indo-West Pacific. This is the first record south the northern archibenthal. of the Tropics and is of particular interest since SOUTH FIJI BASIN

TASMAN BASIN CHATHAM k/ Sf

~ BOUNTY TROUGH

:Jhe'"Snares

Pukakiu Bank

.Auckland Islands KEY

. Gon;ocidor;s umbraculum 'Campbell Island Goniacidar;s parasol

Goniocidar;s" magi

FIGURE 4. The distribution of the echinoid genus Goniocidaris within the New Zealand Region. DAWSON: OCEANOGHAPHY AND MARINE ZOOLOGY 55 fragments of a fossil Lithosoma have been found on the shelf near the Snares. There is a single in the New Zealand Tertiary. record from the Auckland Is. Shelf, and it has also turned up off Macquarie 1. Ceramaster, on the other hand, is a large genus represented in all oceans. The Camp- The only published identification of Henricia bell Plateau species, taken in considerable from l\larquarie is of specimens collected in quantity, has not yet been finally identified Lusitania Bav in 38 fathoms m.A.N.Z.A.R.E. but is not closely related to the only other Sta. 83). These have been identified as Henricia known southern species, C. patagonicus. More obesa (Sladen 1889). This is the first record specimens of Ceramaster have recently been of this species outside the Magellanic- Falklands trawled off Macquarie I., providing another area (Clark 1962, 48). Other Henricia spp. biogeographic link. occur at Kerguelen and Marion Is. in the Kerguelenian Province. These records are further evidence suggesting that Australian Indo-Pacific and cosmopolitan Specimens of H enricia collected from high elements contribute to the fauna of the New spots along the Macquarie Ridge await identifi- Zealand Subantarctic, and that Kerguelenian cation. circum-Subantarctic and Antarctic elements are rare.

HENRICIA CONCLUSION

Henricia is a notoriously difficult genus Since identification and interpretation of the systematically. Despite this, the local repre- recent collections of bottom-living animals of sentatives of this small starfish fronl the New the New Zealand Region is not yet complete Zealand and Subantarctic areas can now be it is premature to generalise about ecological seen to fall into three species with another Henrida and biogeographical relationships within the doubtful species, compacta (Sladen New Zealand Subantarctic, or between this 1889), described from a unique juvenile speci- sector of the circum-Subantarctic region and men taken at "Challenger" station 166 in 275 the Kerguelenian and MageUanic Provinces. fathoms north-west of Cape Farewell. The The faunal links between the interesting distribution of Henricia within the New Zea- Macquarie I. material and New Zealand land Region is shown in Fig. 5. appear much stronger than previously sup- Henricia lukinsi (Farquhar 1898) occurs posed. Reports on the Pycnogonida, the at the Auckland Is., Campbell 1., Chatham nemertean worms, the patellid limpets, the Is. in shallow water, and on the New Zealand brachiopods and the scleractinian and stylas- Shelf, and the Veryan and Pukaki Banks. terine corals are now being prepared. The Although recorded as deep as 300 fathoms, it over-all picture will be considerably clearer seems essentially a shallower water species. with the analysis of these and other important It has now been discovered off Macquarie I. groups and there will probably be as many surprises in their distributions and biogeo- H enricia aucklandica Mortensen, 1925 occurs graphical relationships as have been found on the Chatham Is. shore and is said by Fell amcngst the Echinodermata. (1962c, 38) to "range from Cook Strait south- ward" although substantiated records are not available for the New Zealand Shelf. It is now known from the Campbell Plateau and the Antipodes Is., Auckland Is., and Campbell 1., ACKNOWLEDGEMENTS to depths of over 300 fathoms and appears to be a deeper water species. It has also been I am grateful to Professor H. Barraclough Fell, Dr. taken off Macquarie I. D. L. Pawson, Miss Helen Shearburn-Clark and Mr. D. G. McKnight for identifying the echinoderms Henricia ralphae Fell 1952, is known from referred to in this paper; and to the New Zealand Naval Board for their continued co-operation in making depths to 50 fathoms off South Canterbury and H.M.N.Z.S. Rotoiti and H.M.N.Z.S Endeavour avail- East Otago and is now known to be common able. KEY

Henricio oucklondioe Henricio compocto Henri/:/o lukinsi Henricio obeso Henricil1 rolphl1e Henricio surrll1~o SpecieS: awaiting identification

-- FIGURE 5. The distribution of the asteroid genus Henricia within the New Zealand Region. GODLEY: VEGETATION OF AUCKLAND Is. 57

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NOTES ON THE VEGETATION OF THE AUCKLAND ISLANDS E. J. GODLEY Botany Division, D.S.l.R., Christchurch

The following observations on the vegetation ALTITUDINAL DISTRIBUTION of the Auckland Islands, made between 26 December 1962 and 20 January 1963, are Hooker (1847: 2) made the following impor- additional to or amplify those of Hooker tant note on the vegetation in the north of (1847), Cockayne (1904, 1909) and Moar Auckland 1. "It is especially towards the sum- (1958a, b). mits of these hills that the most striking plants