TANE 29, 1983

CHANGES TO THE MARINE BIOTA OF THE HARBOUR

by F. I. Dromgoole* and B. A. Fostert * Department of Botany, , Private Bag, Auckland t Department of Zoology, University of Auckland, Private Bag, Auckland

SUMMARY

The history of study of the marine biota of Auckland Harbour is briefly reviewed, and it is concluded that there is insufficient documented information to make quantitative assessment of changes that have resulted from reclamation, sedimentation and pollution that have occurred with the development of the Port of Auckland. Losses of mangrove and saltmarsh communities are indisputable, but causes of declines in populations of Zostera, Pomatoceros and Perna are not so clear. On the other hand, a number of species have been introduced, and circumstantial evidence suggests these adventives have arrived as ship- fouling. Cases discussed are Codium fragile tomentosoides, Colpomenia bullosa, Limaria orientalis and Sagartia luciae. The most conspicuous newcomer, the oyster Crassostrea gigas, may have been deliberately introduced.

INTRODUCTION

Regular use of the Auckland Harbour by European ships stems from the early 1800s, so there has been ample opportunity for the introduction of adventive fouling species. Maritime reclamation in Auckland Harbour dates back to about 1860 when shores near the commercial centre were filled and extended as wharves and breakwaters. Modification of habitats and inhabitants of the harbour has now been going on for more than 120 years, but scientific study of them has been of much shorter duration. In this paper we wish to document some additions and alterations to the marine biology. Auckland Harbour has three parts (see Fig. 1). The lower harbour includes southern shores of Rangitoto and Motutapu Islands, and also Motuihe and Browns Islands. The middle harbour is clearly defined; it contains the Port of Auckland and is the most modified. In the middle harbour 45% of the shoreline has been modified by reclamation and 24% of the harbour bed has been claimed for wharves, breakwaters, embankments, causeways and other uses. The upper harbour contains expansive intertidal flats, and is under increasing pressure from surrounding urbanisation. Further local changes are inevitable with the spread of coastal industries and the construction of harbour works,

79 leading to more widespread changes in sedimentation resulting from reclamation and dredging.

Fig. 1. Auckland Harbour (=Waitemata Harbour), subdivisions and localities mentioned in the text. Shoreline modification is shown blacked. (1) , (2) Narrow Neck Beach, (3) Cheltenham Beach, (4) North Head, (5) Torpedo Bay, (6) Stanley Bay, (7) Birkenhead, (8) Westmere Reef, (9) Port of Auckland, (10) Container Terminal, (11) Hobson Bay, (12) Orakei Basin, (13) Okahu Bay, (14) Orakei Point, (15) Mission Bay. HISTORICAL REVIEW

Many early writers of taxonomic accounts of the biota probably used specimens collected at Auckland, but the locality identified was not always specified. Among early Auckland records are those of Hooker (1867) who lists some algae from Auckland, and Hutton (1878) who identified Auckland as a locality for two species of barnacles. The first ecological description of the intertidal biota was that of Oliver (1923). He identified and figured shores at Takapuna and Westmere Reef as examples in his wider account of such communities for all New Zealand shores. Powell's (1937) account of the benthic communities stands as a notable contribution to a part of the harbour's biota that has received very little subsequent study. Studies on the algal flora subsequent to those of Oliver were stimulated with the arrival in 1945 of the first Professor of Botany at 80 the University of Auckland. Professor V. J. Chapman's enthusiasm led to papers on saltmarsh vegetation (Chapman and Ronaldson 1958), intertidal algae (Dellow 1948, 1950, 1953, 1955, Cooper 1951, Carnachan 1952) and subtidal algae (Bergquist 1957, Dromgoole 1965). Work on the marine angiosperm Zostera by Armiger (1965) examined the causes of decline of this species first noted by Hounsell (1935) and Powell (1937). Further interest in the marine biology of the region was rekindled with the arrival of the first Professor of Zoology in 1960. Professor J. E. Morton's book "The New Zealand Sea Shore" (Morton and Miller 1968) contains a number of shore descriptions based on localities in the harbour and makes reference to subsequently published thesis studies concerned with shore ecology (Wood 1962, 1968, Luckens 1976). Fouling species in the harbour were examined by Skerman (1959, 1960b). The pelagic biota of the has been studied by Kramer (1894), and Fuller (1950, 1953), and the relation between gulf and harbour plankton explored by Jillett (1971). In recent years a number of reports have been prepared in association with a variety of environmental proposals, most notable of these is the Ecology Report for the Waitemata Harbour Survey (Chapman and Larcombe 1973). BIOLOGICAL CHANGES

Despite the numerous publications and reports on marine biology that can be identified, replicable quantitative data are rare, and it is difficult to find the exact location of most ecological observations. Only the more extreme biological changes can be deduced, for example, the decline of the Zostera beds (cf. data in Wood 1962) and the now rarity of the green- lipped mussel (Perna canaliculus) on accessible harbour shores. Another example is the decrease in the physiognomic dominance of the tubeworm Pomatoceros cariniferus at Westmere (see Fig. 2, 3). The causes in decline in populations, and resultant community alterations, may or may not be associated with long-term, man-induced causes. Decline of mussels is almost certainly so, because of their edibility and use as fish bait. The causes of changes in populations of eel-grass and tubeworms may be more subtle. The mangrove and saltmarsh flats of Auckland Harbour have been materially modified by reclamation and the construction of roads across them. An example of the associated effects on flora is seen in the changes in Hobson Bay where, prior to 1921, large areas were dominated by the eel-grass Zostera but this has now completely disappeared. Hounsell (1935) suggests that this was due to rapid silting following the construction of the waterfront drive and the railway embankments; Powell (1937) on the other hand states that the bay

81 Fig. 2. Westmere Reef, from Oliver (1922), showing hummocks of the tubeworm Pomatoceros cariniferus. bottom became hard and that shell deposits may have completed the destruction. Armiger (1965) questions whether sedimentation was the only factor involved as epidemic losses of Zostera have been linked to disease. The disappearance of Zostera from the harbour is perhaps the best documented change in the flora of soft-bottom communities apart from the loss of mangroves through reclamation. Hounsell (1935) records the loss of Zostera from Stanley Bay attributing it to an undermining of the formation by the tidal stream following construction of tide deflectors in the harbour. Zostera beds recorded by Armiger (1965) in the Tamaki estuary, Howick Beach, Okahu Bay, Torpedo Bay and Cheltenham have now also disappeared and whilst this can be largely attributed to a disease by the Labyrinthula slime mould (Armiger 1964) it is conceivable that pathogenic susceptibility is enhanced by unfavourable conditions for growth such as increased sediment load, reduced salinity or pollutants. Decline in Zostera abundance is not unique to Auckland Harbour, similar epidemic losses were recorded in the Avon-Heathcote estuary during 1929-1953 and in many Northern Hemisphere locations in the 1930s. Because Zostera beds normally support a rich and varied biota the disappearance may have significant effects on other foodchain linked organisms. Faunal elements associated with the Auckland Zostera beds are listed by Oliver (1923) and Wood (1962) and it seems reasonable to assume that these have also declined drastically in numbers since 1921. Alterations in substrate can also have a marked effect on the local abundance of plants. For example, increased sedimentation reduces the area available for colonisation by the larger perennial brown algae (Carpophyllum spp. ) and the small turf forms (Corallina officinalis) and these may disappear to be replaced by soft shore flora of seasonal species such as Codium fragile subsp. tomentosoides and Gracilaria secundata var. pseudoflagellifera. Pollution effects are not well documented although the discharge of raw sewage was once severe, and still continues on a reduced scale. Wallace and Newman (1953a, b) in a bacteriological survey of beaches to the east of the Orakei outfall, reported serious pollution at Mission Bay with MPN Coliform mounts of 51 per 100 ml and mean diatom counts of 21 per 10 ml. Diatom counts increased with distance from the outfall reaching 286 per 100 ml at Karaka Bay where MPN Coliforms had fallen to 11. They concluded that the southern and perhaps eastern shores of Rangitoto Island were in a semi-permanent state of pollution from the diluted sewage fields. However, Carnachan's (1952) study of benthic algal zonation at Rangitoto Island does not reveal any characteristics that can be attributed to the detrimental effects of this type of pollution. Discharge of sewage at Orakei ceased in 1963—5 but continued on a small scale at North Head and Birkenhead (Jillett 1971) and isolated accidental discharges sometimes together with stormwater

83 run-off have occurred in more recent years. Overseas, improvements in sewage disposal methods have led to dramatic changes in shore ecology (e. g. the Thames estuary in U. K. ) but there appears to be no documented evidence of similar large scale effects in Auckland Harbour since 1965. Changes in the immediate vicinity of the discharge sites might be expected but unfortunately have not been mentioned in any detail. Dellow (1955) found little evidence to indicate that the benthic species compositions at sewage outfalls were markedly different from otherwise comparable sites. ADVENTIVE SPECIES Migration of new species into the New Zealand marine biota has no doubt been going on since the end of the last glaciation but it seems doubtful that climatic change has been responsible for many of the recent ones. There is reason to believe that ship aided dispersal has also operated. Some of this may have operated in days of sailing ships. More recently, shorter and more direct voyages and the changes in New Zealand's trading partners has resulted in a marked change in the last ports of call for vessels arriving in New Zealand, particularly in the period 1950—1980 (Fig. 4). Direct evidence that the adventive species

AUST. TOTAL

1947 50 55 6 0 65 70 7 5 79

U. K.

1947 50 55 60 6 5 70 75 79 JAPAN

U. S. A.

nil

Fig. 4. Auckland Harbour shipping movements 1947-79. The bars indicate the numbers of vessels whose first New Zealand port of call was Auckland and whose previous port was in the country indicated. The major change in this period has been the marked increase in the numbers arriving directly from Japanese ports.

84 have been transported as ship-fouling or as dunnage (e. g. packing around shellfish or other seafood on fishing vessels) is difficult to obtain and in many cases it is not possible to determine an exact date of arrival. Table 1 lists those species which are regarded as adventives to Auckland Harbour.

Table 1. List of adventive species which now occur in Auckland Harbour or adjacent localities. Dates of introduction are estimates.

SPECIES GROUP DATE EARLIER REFERENCE OF LOCALITIES INTRO• DUCT• ION Watersipora Bryozoa 1957 Australia Skerman 1960a arcuata Ban ta Ciona Ascidiacea 1960 Northern Morton & Miller intestinalis hemisphere 1968 (Linnaeus) Crassostrea 1964 Japan, Australia Dinamani 1971 gigas (Thurnberg) Limaria Bivalvia 1972 Japan - Australia Grange 1974 orientalis (Adams & Reeve) Codium fragile Chlorophyceae 1973 Japan Dromgoole 1975 (Sur) Hariot subsp. tomentosoides (Van Goor) Silva Theora lubrica Bivalvia 1972 Japan Climo 1976 Gould Hydroclathrus Chlorophyceae 1974 Kermadec Is Johnson & Dromgoole clathratus 1977 (C Ag) Howe Sagartia luciae Anthozoa 1977 Japan - Panama New record Verril Pyromaia Brachyura 1978 Japan Webber & Wear 1981 tuberculata (Lockington) Colpomenia Phaeophyceae 1980 Japan Parsons 1982 bullosa (Saunders) Yamada

It is inevitable that not all adventives have been detected. For one thing, early documentation of the biota may not have accounted for some 'rare' (or rarely observed) species. Indeed, some marine taxa, e. g.

85 amphipods and copepods, have still not been adequately studied by biologists in Auckland, so that it is hard to know what is new or indigenous within these groups.

Codium fragile (Sur) Hariot subsp. Tomentosoides (Van Goor) Silva. Fig. 6. This siphonous green alga, apparently, a native of Japan, has been recorded as a vigorous adventive to many parts of the Northern Hemisphere. First recorded in New Zealand at the eastern wall of the Container Terminal in Auckland Harbour in 1973 (Dromgoole 1975), the plants have spread to and established in many localities (Dromgoole 1979). The adventive species is now frequently found in the seaweed drift on Auckland beaches as intact plants attached to a substratum ballast. This alga is now a seasonally (November to April) frequent component of lower littoral and sublittoral communities on rocky and soft shores and a seasonal dominant of fouling on wharf piles, pontoons and buoys. Introduction is tentatively attributed to shipping as the plants have a perennial basal system which becomes firmly attached to fouling surfaces. The primary method of reproduction is by asexual swarmers but vegetative propagation from fragments is also possible and may have contributed to the rapid spread. The plants consist of one to several erect fronds abundantly branched in a dichotomous or fastigiata fashion and attached to the substratum by a spongy basal disc. In general morphology the plants are very similar to the native C. fragile subsp. novae-zelandiae (not recorded in Auckland Harbour but occasionally found on the West Coast beaches of Piha and Bethells) but the adventive can be distinguished on microscopic examination by the broad median or sub-median construction and sharp mucron of utricles taken from about 2 cm below the branch tip. In contrast utricles taken from an equivalent position from subsp novae-zelandiae have a cylindrical or slightly elevated shape with bluntly mucronate tips. These differences are indicated in Fig. 5. There are no detailed studies of the interaction of this adventive species with endemic organisms, but the plants are rarely dense enough to physically displace or blanket other organisms and thus may be considered a useful addition to the harbour ecosystem as primary producers. However, mature plants can be as much as 2-3 kg fresh weight and their large size could pose a hydrodynamic problem by fouling of pontoons and buoys, or interfere with mariculture enterprises. How this plant was transported to New Zealand is not certain. Movement in the Northern Hemisphere has been speculated to occur by accidental transportation with seed oysters or as ship fouling. It is unlikely that the Auckland plants were inadvertently imported with seed oysters (Mr D. Waugh pers. comm. ). Considering the probable date of introduction (1974), and the site of the first recorded population, it is

86 Fig. 5. Typical utricle form in (a) Codium fragile subsp. novaezelandiae (J. Ag) Silva and in (b) Codium fragile subsp. tomentosides (Van Goor) Silva. Utricle form in Codium becomes distorted as the branch develops and diagrams refer to standard samples taken at a distance of 2 cm from the tip. speculated that the plants may have been introduced as ship-fouling, probably from Japan.

Hydroclathrus clathratus (C Ag) Howe. Fig. 7 This attractive perforated brown alga is cosmopolitan in tropical and warm temperate waters. It was previously recorded in New Zealand waters only as fragments from the Kermadec Islands but has recently been found at several localities on the east coast of Northland (Johnson and Dromgoole 1977). Hydroclathrus is probably a recent adventive. It was not reported in the extensive floristic studies of Lindauer between 1935 and 1953 (Cassie 1971), nor was it noted at Whangarei sites under surveillance since 1960 (Professor J. E. Morton pers, comm. ) until 1974. Plants are now found infrequently in sheltered harbour sites on rocks or cobbles in the lower intertidal or subtidal (down to 2 m below low water) region of shallow bays usually associated with the dominants Colpomenia sinuosa and Corallina officinalis. It seems unlikely that this plant was introduced by shipping as it is not noted as a fouling species nor has it a strongly developed anchorage system. Gas bubbles trapped within the thallus of young imperforate plants would provide sufficient

87 Fig. 6. Codium fragile tomentosoides (Van Fig. 8. Limaria orientalis (Adams & Reeve), Goor) Silva, from Narrow Neck, 1975. from Westmere Reef, 1982. buoyancy and this mechanism may have permitted the recent southward extension of the range as surface drift in the warmer waters of the North Auckland current - an algal parallel to the occurrence of tropical and subtropical fish (Russell and Ayling 1976) and molluscs (Powell 1976) that have invaded northern New Zealand waters in recent years.

Colpomenia bullosa (Saunders) Yamada This brown alga was first recorded in New Zealand by Parsons (1982) from collections made at Leigh in 1980. Previously known from northern Japan, and from Alaska to central California, it has been suggested by Parsons that this plant may have arrived in New Zealand associated with ships. A closely related species Colpomenia peregrina Sauv. is recognised as an adventive in the Northern Hemisphere and details of its spread into Europe and northwards to United Kingdom are outlined in Jones (1974). However, this species is probably endemic to New Zealand, together with the common Colpomenia sinuosa (Roth) Derb. et Sol, as both are found in the early herbarium collections of Tilden and Lindauer.

Limaria orientalis (Adams and Reeve) Fig. 8 This bivalve became noticeable in the early 1970s (Grange, 1974), and it is now a common constituent of low tide and benthic faunas of the harbour where coarse shell and rubble substrata occur. In 1980 it occurred in grab samples in the Motutapu-Motuihe Channel in densities up to 100 per m2. Lists of species in these channel samples do not differ from the faunal lists compiled by Powell (1937) except for the occurrence of two species, L. orientalis, and a small spider crab Pyromaia tuberculata. The crab was first found at Orere Point in 1978 by Dr Roger Grace and a full assessment of it is being prepared by him and Dr J. Yaldwyn. L. orientalis has become important in the ecology of the area. Snapper populations have been sampled off Waiheke for the years 1979—81 for class analysis in a University of Auckland Stage III ecology course. From gut content analyses (Fig. 10), it is clear that Limaria has become an important part of snappers' diet; over 50% of fish from 4 age classes were found to have the bivalve in their guts, and by volume these become dominant dietary components in the larger fish. Its present colonisation of Auckland east coast waters may not be the first in the history of New Zealand. L. orientalis is known as a Miocene and Pleistocene fossil (Powell 1979). Whereas the present populations could have been introduced by shipping, prehistoric introductions may indicate the possibility of larval transport (or rafting of adults in floating debris) or reflect climatic change.

89 10-20 cm 20-30 cm

(n=12) (n=13) Limaria

Hermit Crabs

CH Crabs

HU Maoricolpus

Amphipods 30-40 cm Fish (n=13)

Fig. 10. Proportionate representation of food items in the stomachs of 51 snapper {Chrysophyrus auratus), separated into 4 size groups (standard lengths), caught off Waiheke in July 1981.

A new anemone for Auckland Harbour Fig. 9 Since 1977, a foreign anemone has been noted occurring in Orakei Basin, and has been more recently found at other localities such as Bucklands Beach and Westmere Reef. It is never in very large numbers, and occurs intertidally in muddy and somewhat brackish habitats. Specimens were identified by Dr Cadet Hand as 'Haliplanella luciae which is the name currently applied to a common Northern Hemisphere fouling sea anemone once known as Sagartia luciae. A full synonymy is given by Hand (1955). Apparently, however, the has not been resolved; for one thing Hand's genus Haliplanella is preoccupied by a genus of polychaetes, and at least 2 other earlier species names occur in the literature for similar sagartids, namely pus tutata for specimens described in 1887 from Beaufort, North Carolina, and lineata for specimens described in 1870 from Hong Kong. Dr Hand is reviewing the position of this species, or species-complex, and the exact identity of the Auckland specimens awaits his pronouncement. Whatever its name, the anemone is distinctive among Auckland species by its bush of light green tentacles arising from a dark green 90 column with vertical orange stripes. Like the brown-tentacled Diadumene novaezelandicus, it exudes acontia through cinclides in the column wall when provoked. Maximum height of Auckland specimens is about 15 mm. Hand (1955) concludes that the north Pacific is the original home of this anemone, probably the north-western Pacific. It is assumed to have subsequently been spread to the Pacific coast of North America, the Suez Canal, Texas, New Haven (by 1892), England (by 1896), and later in Europe. It has not previously been recorded from the Southern Hemisphere although Dr Hand (pers, comm. ) reports specimens have recently been collected from Brazil. Ricketts and Calvin (1939) note that on the Pacific coast of North America, the anemone has appeared at or near places where Pacific oysters (Crassostrea gigas) are grown. Its dispersal by the transport of oysters, and by fouled ships, seems indisputable. On the Asian coast, the anemone reproduces sexually (Uchida 1932) but North American populations seem to be single-sexed clones (Hand 1955).

Crassostrea gigas (Thurnberg) Fig. 11 C. gigas is the Pacific oyster, sometimes called the Japanese oyster, that is used extensively around the world in mariculture. The species is now well established on harbour shores in northern New Zealand. It was first reported from off oyster-farm racks in the Mahurangi Harbour in 1970 (Dinamani 1971) and has since extended its range south to the Marlborough Sounds and Tasman Bay (Dinanami 1981, Bull 1981). The mode of arrival of this oyster in New Zealand is obscure, but it is clear that it was established by the early 1960s. A photograph of oysters in a thesis by Rainer (1965) shows a Pacific oyster from the Kaipara Harbour, and Mr Les Curtin reports the existence of a shell collected from Whangarei in 1958 serving as an ashtray on the desk of an employee of the oyster section of the Ministry of Agriculture and Fisheries. The coincidence of these places with early attempts at commercial oyster 'farming' is suggestive of deliberate introduction of the species to New Zealand to aid 'the industry'. This was certainly done in Australia (Thompson 1952). It was not until 1978 that numbers of C. gigas in the Mahurangi Harbour exceeded catch of the native rock oyster (Saccostrea glomerata) on spat-collecting sticks. Also in the late 1970s, C. gigas became noticeable on Auckland Harbour shores, and it is now well established and competes with the native oyster; its faster growth rate results in smothering of the spat of the native species. On Westmere Reef, the black basalt of the intertidal is now conspicuously encrusted with C. gigas (Fig. 3), contrasting with the situation in the 1920s (Fig. 2) when the tubeworm Pomatoceros cariniferus formed a white zone at slightly lower levels. Of all the recent marine invertebrate adventives the Pacific oyster is

91 Fig. 11. Crassostrea gigas (a-c) and Saccostrea glomerata (d-f): upper view of upper (right) valve (a, d), inner view of upper valve (b, e), inner view of lower (left) valve (c, f). Drawn to emphasise the more finely frilled growth ridges in C. gigas and dentations on dorsal and ventral articular edges near the umbo of both valves in S. glomerata. the most obvious; its tendency towards massive settlements can reinforce the oyster zone on rocky shores. Its subsequent fast growth rate makes it a better commercial proposition to oyster farmers than the native species. Whereas once it was regarded as a pest, it is now a marketed species. The difficulties in distinguishing the two species stems from the variety of shell forms of both species. Large shells (over 5 cm long) with sharp frilly and raised growth ridges will invariably be C. gigas. Eroded shells of both species are externally similar. Empty shells of S. glomerata have a series of ridges on the margin near the hinge that produce a grating noise when scratched. These ridges are absent in C. gigas. Distinguishing features of these two taxa are shown in Fig. 11. In

92 S. glomerata, colour is purplish-black on white whereas in C. gigas shells are dark to light brown on white. The ecology of the two intertidal species are not exactly the same; S. glomerata can occur higher on the shore than C. gigas because it is tolerant of emersion conditions. Protected low tide and oyster rack populations possibly provide the breeding stock of C. gigas. Monitoring of the relative dominance of these two species at a number of Auckland localities would be a useful contribution to our understanding of the ecology of the adventive.

ON THE SOURCE OF INTRODUCTION

Some exotic introductions into the northern New Zealand marine biota, such as the crab Scylla serrata and the cone shell Conus kermadecensis, are conceivably by dispersal of long-lived larvae. Both of these cases, involving only few adults being found, are in coastal regions which are more subject to currents from the north-west which could carry the larvae from lower latitudes. Their introductions have not been followed up by the establishment of breeding populations. But many post-Pleistocene invasions must have been successful, and Hydroclathrus clathratus seems to be a very recent instance. The establishment of new species in ports points more towards introductions involving transport by ships, pontoons, etc. Prior to 1956 overseas shipping entering New Zealand ports was predominantly Australian and this trans-Tasman traffic has been implicated in the transport of Watersipora cucullata (Skerman 1960a). In the Port of Auckland some other generally acknowledged fouling introductions have been the ascidian Ciona intestinalis (Morton and Miller 1968), the polychaete tubeworm Hydroides norvegica (Allen 1953) and the opisthobranch Thecacera pennigera (Willan 1976). Marine species that make up marine fouling and that have widespread distributions elsewhere in the world, could also be long-established and ship-dispersed long before recording of marine faunas commenced. Barnacles are well-known for being spread by shipping. Fouling barnacles in the Waitemata Harbour are Elminius modestus (which has been spread to overseas localities from New Zealand), and 3 species of Balanus of less certain native status - Balanus trigonus, B. variegatus and B. amphitrite. All of these three species may have been brought to northern New Zealand waters by shipping. B. amphitrite has a particularly restricted distribution. It occurs abundantly only in the modified sea conditions of the Orakei Basin. In recent years there has been a dramatic increase in the number of vessels entering New Zealand whose last post of clearance was Japan (see Fig. 4) and it may be more than a coincidence that many of the recent arrivals in the Port of Auckland are common in the waters of

93 Japan e. g. Codium fragile subsp. tomentosoides, Limaria orientalis, Crassostrea gigas, the new anemone, Thecacera pennigera and Pyromaia tuberculata.

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