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Invertebrates, P 34-44Okak006.Pdf guilds of species have been recognised (Russell 1971a). Hence, there are 12 guilds altogether, comprising bottom grubbers, bottom cleaners, bottom fossickers, invertebrate grazers, plant grazers, plant browsers, bottom stalkers, invertebrate browsers, general predators/scavengers, parasite cleaners, plankton pickers and midwater hunters. Virtually every phylum of animals and the four major groups of macroscopic algae are utilised as food items. Using a relatively simple index of utilisation, based upon the intensity of consumption of food items and the dispersion of that food among the predators, Russell (1971a) determined that the 10 highest values were scored, in decreasing order, by amphipods, brachyurans, fishes, gastropods, copepods, errant polychaetes, anomurans, bivalves, rhodophytes and ecninoids. Additional information on food and feeding is provided by Ayling (1968b), Cassie (1956a), Coleman (1972), Doak (1972), Godfriaux (1969, 1970a,b) and Russell (1971b, 1975). D. Behaviour Apart from a partial study of the behaviour of blennies (Anderson 1973), the cleaning behaviour of wrasses (Ayling & Grace 1971), and the feeding behaviour of leatherjackets (Ayling 1976a), most information is scattered in ecological works. Ayling (1968) and Russell (1971a), for example, provide a considerable amount of data on the way fish behave when seeking out and ingesting food, meanwhile commenting on whether they are schooling (see earlier) or solitary, and whether or not territoriality is exhibited. Some fish have definite homes (hiwihiwi, blue cod, red cod) while others have limited ranges (banded sea perch, red moki, black angelfish, scarlet parrotfish and possibly the banded parrotfish and twister). Even school-forming koheru, when juvenile, appear to have a limited home range. A school of 200-300 occurred over an artificial reef off Goat Island for several months. Territoriality is marked in three species of Tripterygion (T. bucknilli, T. varium, T. capito), especially during the breeding season. Hiwihiwi and red mullet appear to form family groupings while the crested blenny has a definite social hierarchy. Several species are nocturnal in their habits (grandfather hapuka, yellow moray eel, conger eel, red cod, bastard red cod, roughie and big-eye) while a few (drummer, parore) are largely crepuscular. The giant boarfish, Sandager's parrotfish and Pseudolabrus miles are sexually dimorphic and the butterfish exhibits cryptic behaviour and colouration. P. miles and Sandager's parrotfish are facultative cleaners (Ayling & Grace 1971). Dellichthys morelandi lives in a commensal relationship with Evechinus. INVERTEBRATE BIOLOGY The invertebrate life of the Marine Reserve appears visually fairly diverse, although, at the time of this writing, only 942 species have been definitely recorded within the Reserve boundaries. Much of the research carried out at the Leigh Laboratory has been on aspects on invertebrate biology, with some emphasis on the Mollusca, partially reflecting the numerical preponderance of the molluscs (299 species) and no doubt also of the malacologists. Much 34 information is found in general works on community biology. The literature is mentioned below under each major invertebrate group. Where an authority is not quoted for a record it is to be assumed that the source is the Faunal Index of the Leigh Laboratory. A. Protozoa. The protozoa are underrepresented in faunal lists (51 species total). About 40 foraminiferans (Eade 1967; Thompson 1975) and some conspicuous ciliates (folliculinids and Zoothamnion) are reported. Most of the foraminifera were recorded from a sample of terrigenous sand, taken at 24m off Goat Island. The total population was estimated at 10 individuals per gram of sediment, one fifth of which were "living" (i.e. responding to staining techniques, the results of which tend to be equivocal). Of the 38 species from the one sample, 18% were porcellanous, 52% were hyaline-calcareous and 29% arenaceous. The ratio of planktonic to benthonic forms was 0.09 (Thompson 1975). These 38 species probably represent about half of the forams that may be expected in the sediments of the Reserve (pers. comm. Dr M.R. Gregory, Geology Department, University of Auckland, 1976). Future records may include species of particular interest to specialists. For example, a common foraminiferan at Goat Island Bay, which settles readily on objects in aquaria, is Dendronina arborescens. This species, originally described as a new genus and species from 140-600m off North Cape and Three Kings in 1922, was not again recorded until its rediscovery at Goat Island Bay in 1967 (Mr J.V. Eade, N.Z.O.I., in litt., 1968). Apart from foraminifera, free-swimming protozoans are numerically abundant, interstitially in sediments and in the seawater system of the Leigh Laboratory, but none of these has been identified. B. Porifera. The sponges have been fairly well characterised taxonomically (81 species) (Hogg 1967, Bergquist 1961, 1968, 1970) and Goat Island Bay is the type locality of one species (Timea aurantiaca). Hogg gave ecological data for 33 species with notes on the breeding seasons of 9 species. Sponges are significant components of subtidal communities (Ayling 1968, 1974d). On the northeast point of Goat Island Ayling (1968) has estimated the standing crop of sponges at varying depths on different slopes (table 6). Studies on the aggregation of dissociated sponge cells were carried out by Reid (1969). It was found that monospecific aggregates differentiated rapidly into a form resembling an adult sponge, whereas even forced bispecific aggregates (viz. Halichondria spp.) did not. C. Mesozoa. Mesozoans have not been recorded from the Reserve, but Octopus and Robsonella, which occur locally, are known hosts of five species of dicyemid mesozoans at Kaikoura (Short 1969; Short & Hochberg 1971), and it is possible that, if they were examined, they may be found to contain these parasites. D. Cnidaria. Cnidarians are abundant numerically, especially during seasonal peaks of hydroids and jellyfish, but are not diverse taxonomically. Hydroids have not been well characterised and only 24 species of Hydrozoa are known within the Reserve. Ayling (1968b) has noted the zonation with depth of 8 species of hydroid off the northeast point of Goat Island. 35 Jellyfish (Aurelia aurita, Phyllorhiza punctata, Cyanea capillata) occur seasonally, and Aurelia washes ashore at times in great numbers. The Anthozoa (17 species) are represented by the orders Alcyonaria, Stolonifera, Actiniaria, Corallimorpharia and Madreporaria. Two species of coral are found commonly (Flabellum rubrum, Culicia rubeola) and Russell (197la:64) records discovering Paracyathus conceptus on his artificial reef off the north-east point of Goat Island. Squires (1963) formally recorded F. rubrum from Cape Rodney. The anemone Actinothoe albocincta is common subtidally, attaining 36.5 g/m2 dry weight (representing 12.41% of the biomass)at 13m depth on flat rocky areas of Goat Island, where it may occur in numbers >1000/m2 (Ayling 1968b, 1974b). Light intensity and water movement affect the distribution of Actinothoe, which occurs on vertical surfaces at 6m and on flat surfaces at 13-16m (ibid). Depth m Slope % community grams/m2 dry weight 6 0° 13 73.0 12 0° 11 31.2 12 90° 97 700.0 12 120° 55 234.0 17 60° 31 151.0 20 0° 55 184.0 J L Table 6: Percentage contribution of sponges to the standing crop of the biota of the north-east point of Goat Island. (From data of Ayling 1968.) Depth m Carnivores Deposit feeders Filter feeders 6 2.2 0 0 12 8.0 5.4 3.3 17 4.5 1.3 0.5 20 0 3.4 0 Table 7: Standing crop, in grams per m2, of polychaete worms on the north-east point of Goat Island. Each figure is the total for that depth over all slopes. (From data of Ayling 1968.) 36 E. Ctenophora. Pleurobrachia pileus and a species of Bolinopsis occur in the plankton while a creeping species, Coeloplana willeyi, is a benthic form. First recorded from Pakiri (Gordon 1969), this species has since been discovered on Pecten shells 19m off Goat Island. These are the only known New Zealand localities of this wide-spread Indo-Pacific species. An unidentified cestid ('Welamen sp.) occurs in outer Gulf waters which might possibly be found in the Reserve. F. Platyhelminthes. Parasitic flatworms (trematodes and cestodes) occur in fishes, birds, cetaceans and some invertebrates in Reserve waters, but none has been formally recorded. Of the larger poly clad Turbellaria, 12 species have been found (by Prof. J.J. Holleman, Merritt College, California) but not all have been described. The biology of none of the local species is known. Many small acoels and rhabdocoels occur in algal or invertebrate biotopes but these have not been studied by any worker. G. Gnathostomulida. These inhabitants of anoxic marine sediments have not been discovered in New Zealand. H. Nemertea. Several nemerteans occur intertidally (Morton & Chapman 1968; Morton & Miller 1968) but none has been identified to species level. I. Acanthocephala. None of these parasites has been formally recorded from Reserve waters, but it is likely that some fish and sea birds may harbour them. J. Entoprocta. Pedicellina hispida and Barentsia gracilis are frequently found in association with bushy bryozoans or at the bases of some small low-tidal algae. An unidentified loxosomatid occurs at times on walls of aquaria at the Leigh Laboratory. K. Aschelminthes. Free-living nematodes are abundant in algal and invertebrate biotopes and in sediments but little information exists on nematodes in local waters (Ditlevsen 1929). Parasites have been found in snapper and other fish (by students at the Leigh Laboratory) but not identified. Much basic taxonomic work is needed on both parasitic and free-living forms. A rotifer, attached to the body wall of a holothurian (possibly Kolostoneura novaezelandiae) from the Echinoderm Platform has been recorded. Gastrotrichs, kinorhynchs and nemato- morphs are not known from Reserve waters. L. Annelida. The polychaetes of the Reserve have been well characterised (Whitley 1966). Of the Errantia 26 species are known, with 30 species of Sedentaria.
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