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              <editor role="editor" TEIform="editor"><name type="person" key="name-102052" TEIform="name">J. W. Dawson</name></editor>
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<titlePart type="main" TEIform="titlePart"><hi rend="c" TEIform="hi">Tuatara</hi></titlePart>
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<docImprint TEIform="docImprint"><hi rend="c" TEIform="hi">Journal of the Biological Society<lb TEIform="lb"/>
Victoria University of Wellington<lb TEIform="lb"/>
New Zealand</hi><lb TEIform="lb"/>
<hi rend="c" TEIform="hi">Volume</hi> 22 <hi rend="c" TEIform="hi">Part</hi> 3 <hi rend="c" TEIform="hi">February</hi> 1977</docImprint>
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<head TEIform="head"><title level="a" TEIform="title"><hi rend="c" TEIform="hi">A Brief History of New Zealand Marine Biology</hi></title></head>
<byline TEIform="byline">by <name type="person" key="name-170577" TEIform="name">George Putnam</name><lb TEIform="lb"/>
Harvard University</byline>
<div2 id="t1-body-d1-d1" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">The Beginnings</hi></head>
<p TEIform="p">Long before the first European explorers caught sight of New Zealand, the seas around the country were being examined and exploited by the Maori inhabitants. When Sir <name type="person" key="name-123818" TEIform="name">Joseph Banks</name> first arrived, he found proof of this in the form of ‘vast heaps of shells, often many waggon loads together, some appearing to be very old’ (Morrell, 1958). He also found tools and ornaments made from the bones and teeth of fish, seals, and even whales. More recent archaeological evidence indicates that the Maoris included in their diet virtually all the types of fish found today in New Zealand coastal waters, plus more than twenty-five species of molluscs, and assorted sea urchins, crayfish, and even ascidians (Hamilton, 1908).</p>
<p TEIform="p">The Maoris developed their own theories of oceanography and marine biology. Some of these were fanciful, such as their belief that the tides were caused by a huge sea monster named Panata, but others were based on somewhat ‘scientific’ observations. For example, after observing the density of fish around upwellings and fresh water springs near the coast, they speculated that all fish were produced from fountains in the sea called Rangiriri. The Maoris also developed special lunar calendars which indicated the best times to catch certain fish (Hamilton, 1908).</p>
<p TEIform="p">The first European to visit New Zealand, Abel Jantzoon Tasman, apparently did little to introduce western methods of scientific investigation. He sailed along the western coast of New Zealand for nearly a month in 1642-43, but he made only two brief landings. There is no mention in his journals of any natural history observations (Hocken, 1896).</p>
<p TEIform="p">After Tasman's departure, more than 125 years were to pass before another European expedition came to New Zealand. The arrival of Captain <name type="person" key="name-207700" TEIform="name">James Cook</name> in 1769 marked the beginning of the scientific investigation of the seas of the region. Cook had been sent out,
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sponsored jointly by the British Admiralty and the Royal Society of London, to Tahiti to observe the transit of Venus in 1769, He was accompanied by two capable scientists, <name type="person" key="name-123818" TEIform="name">Joseph Banks</name> and Daniel Carl Solander, the latter having been a pupil of Linnaeus. Moreover, Banks spent £10,000 of his own money to equip the vessel for scientific research.</p>
<p TEIform="p">On his journey homeward from Tahiti, Cook spent from October 1769 through March 1770 circumnavigating both the North and South Islands. During this time, Banks and Solander examined many marine organisms collected by diverse methods including handlines, ‘seans’, ‘druge’, and ‘dipping net’ (Morrell, 1958). They took numerous specimens of fish, invertebrates, and seaweeds, most of which had never before been recorded. Unfortunately, the scientific results of the voyage were never completely and systematically published. An official narrative of the expedition compiled from Cook's and Banks's journals by John Hawkesworth was published in 1773, but Cook's journal itself was not printed until 1893, and Banks's journal remained unpublished until it appeared in a considerably edited and modified version in 1896.</p>
<p TEIform="p">Cook returned to New Zealand in 1773 and 1774 during the course of his second voyage, an expedition to explore the Antarctic seas, This time he was accompanied by the father and son naturalist team of J. R. and <name type="person" key="name-123817" TEIform="name">George Forster</name>. (Banks had originally planned to serve as naturalist of this voyage, but he withdrew after an argument with the Admiralty.) During the voyage, New Zealand was visited three different times: from March to June 1773 at Dusky Bay and Queen Charlotte Sound; November 1773 at Queen Charlotte Sound; and October to November 1774, also at Queen Charlotte Sound.</p>
<p TEIform="p">The Forsters collected many marine organisms, but, as with Cook's first voyage, very few of the results were published until many years after the expedition returned. An official journal of the voyage written by Cook was published in 1777, and generalised accounts by George and <name type="person" key="name-124833" TEIform="name">J. R. Forster</name> appeared in 1777 and 1778 respectively, but none of these contained much scientific information. Some of <name type="person" key="name-124833" TEIform="name">J. R. Forster</name>'s descriptions of New Zealand fish were used in Bloch's <hi rend="i" TEIform="hi">Systema Ichthyologiae,</hi> 1801, but the complete systematic account of the scientific results of the voyage, written by <name type="person" key="name-124833" TEIform="name">J. R. Forster</name>, was not published until 1844. By that time Forster was dead, and many of his specimens had been described and renamed by other workers.</p>
<p TEIform="p">In February 1777, while on a voyage to search for a northern passage through the North American continent, Cook made yet another visit to New Zealand. This time he was accompanied by <name type="person" key="name-121726" TEIform="name">W. Anderson</name> as naturalist, but it seems that little biological work of any sort was carried out.</p>
<p TEIform="p">Despite the various problems surrounding the publication of the scientific results, Cook's voyages provided a solid beginning for the science of marine biology in New Zealand. During the course of
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the three voyages, about 40 species of molluscs, plus many other invertebrates, were taken from these waters. Cook's naturalists also identified 65 species of New Zealand fish, and their work proved to be so thorough, relative to the other naturalists of the period, that over the next 65 years only 25 more new species of fish were found in the area (Richardson, 1843).</p>
<p TEIform="p">At the same time as Cook was exploring the South Pacific, two French expeditions came to New Zealand. In late 1769 J. F. M. de Surville stopped on the North Island, while on the way to Tahiti, to allow his crew to recover from scurvy. He made a few comments on the anti-scorbutic plants found in the area, but there is little in his narrative to interest the marine biologist (McNab, 1914). Three years later de Surville's countryman, Marion de Fresne, spent about three months in New Zealand on a private journey to Tahiti, reputedly ‘in the cause of science’. De Fresne and his second in command, Crozet (who took command when de Fresne was killed by Maoris), made a few comments on whales, porpoises, and fish. Their most interesting remarks from a marine biological point of view came from their observations of the migration of fish along the coast and of the relative abundance of fish in Cook Strait (Fell, <hi rend="i" TEIform="hi">et al.</hi>, 1953).</p>
<p TEIform="p">After Cook left New Zealand for the last time, a series of less well known expeditions came to these shores and made small contributions to the knowledge of the marine biology of the area. In 1791 Capt. George Vancouver, who had served under Cook, put into Dusky Sound for provisions on his way to explore the north-west coast of North America. He was accompanied by a competent naturalist, Archibald Menzies, but Menzies' marine observations seem to have been limited to a few comments on sea birds and seals (McNab, 1907). Vancouver was followed in 1793 by a Spanish fleet under Malaspina which visited Doubtful Sound and in 1820 by a Russian expedition commanded by Thaddaus Bellinghausen which stopped at Macquarie Island. Very little is known about Malaspina's observations because it appears that no journal of his voyage was ever published. Bellinghausen did publish some descriptions of the appearance and behaviour of penguins and ‘sea-elephants’ of Macquarie (McNab, 1907).</p>
<p TEIform="p">Around the beginning of the nineteenth century, a new source of information appeared in the form of whaling and sealing expeditions. The first whaling ship to visit New Zealand waters arrived in 1791, and by 1802 at least seven boats were reported to be whaling in the area. Similarly, the first sealing gang landed in Dusky Sound in 1792, and it was soon followed by many others. These expeditions were primarily interested in economic success, of course, but they did make scientific contributions. The British naturalist, J. E. Gray, noted that the French whalers who visited New Zealand ‘were constantly sending zoological specimens to Paris’ (Dieffenbach, 1843). Unfortunately, the whalers and sealers rarely published any of their
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biological findings themselves, but some of their work did appear through other channels. For instance, Bellinghausen included information from sealers in the journals of his voyage, and J. A. R. Blosseville, a Frenchman, used data from whaling expeditions in his <hi rend="i" TEIform="hi">Memoire Geographique sur la Nouvelle Zelande</hi> published in 1826 (McNab, 1907).</p>
<p TEIform="p">Perhaps the one ship that contributed the most to New Zealand marine biology in this period was a French corvette, originally named <hi rend="i" TEIform="hi">Coquille</hi>, which visited the islands three times between 1824 and 1840. She stopped in at the Bay of Islands in April 1824 on a voyage of scientific discovery around the world under the command of L. I. Duperry with Dumont D'Urville as second officer. R. P. Lesson was the naturalist on board, and, assisted by the ship's surgeon, P. Garnot, he described several species of fish and molluscs from New Zealand. Included among these were three new species of fish and nine new species of molluscs. These descriptions, with some illustrations, were published in an account of the zoology of the voyage in 1826-32.</p>
<p TEIform="p">In 1827 the <hi rend="i" TEIform="hi">Coquille</hi>, now renamed <hi rend="i" TEIform="hi">Astrolabe</hi>, returned to New Zealand under the command of Dumont D'Urville in the course of another ‘extended voyage of discovery and exploration’ for the French government (Wright, 1950). This time the naturalists were J. R. C. Quoy and J. P. Gaimard. They appear to have been primarily interested in molluscs, and in their official reports of the voyage, published in 1833, they described 85 different species from that group. Their work was sufficiently important to prompt <name type="person" key="name-209371" TEIform="name">H. Suter</name>, a prominent twentieth century mollusc expert (see below), to call it, ‘The most valuable contribution to New Zealand conchology in the early days’ (Suter, 1913). In addition to their observations on molluscs, Quoy and Gaimard also described many species of tunicates. radiates, and cirripedes.</p>
<p TEIform="p">The <hi rend="i" TEIform="hi">Astrolabe</hi> made her third and final stop in New Zealand in 1840 on a voyage to the Antarctic, this time accompanied by another corvette, <hi rend="i" TEIform="hi">Zelee.</hi> D'Urville was again in command, and his naturalists were J. B. Hombron and H. Jacquinot. They described and illustrated 15 species of molluscs from New Zealand waters, plus several species of fish and crustacea (Fell <hi rend="i" TEIform="hi">et al.</hi>, 1953).</p>
<p TEIform="p">During the period from 1835 to 1850, four of the most famous naturalists of the nineteenth century visited New Zealand. <name type="person" key="name-121361" TEIform="name">Charles Darwin</name>, perhaps the best known biologist in history, arrived in 1835, and he was shortly followed by his illustrious countrymen, <name type="person" key="name-208267" TEIform="name">J. D. Hooker</name> and T. H. Huxley. Also during this period, a leading American naturalist, James D. Dana, visited the islands. Although their visits to New Zealand did not appear to significantly affect their respective future careers (a fact that dismays many chauvinistic Kiwis), these men did contribute to the knowledge of the marine biology of the area.</p>
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<p TEIform="p">Darwin visited the Bay of Islands for nine days in December 1835 while serving as naturalist on <hi rend="i" TEIform="hi">H.M.S. Beagle</hi> on a surveying voyage around the world. In the official record of the voyage, published in 1842, he described a few species of fish found in New Zealand, but his main contributions to marine biology came from his later works including those on coral reefs (1851), cirripedes (1851, 1854), and, of course, evolution (1859). Perhaps Darwin's lack of collecting in New Zealand could have been caused by his feelings about the country, for on his departure he said. ‘I believe we were all glad to leave New Zealand. It is not a pleasant place’ (Darwin, 1959).</p>
<p TEIform="p">Hooker accompanied James Clark Ross on the vessels <hi rend="i" TEIform="hi">Erebus</hi> and <hi rend="i" TEIform="hi">Terror</hi> on a voyage ‘for the purpose of investigating the phenomena of Terrestrial magnetism in various remote countries, and for prosecuting Maritime Geographical discovery in the high southern latitudes’ (Fell <hi rend="i" TEIform="hi">et al.</hi>, 1953). The <hi rend="i" TEIform="hi">Erebus</hi> and <hi rend="i" TEIform="hi">Terror</hi> paid two visits to New Zealand: at the Auckland and Campbell Islands in November to December 1840, and at the Bay of Islands in August to November 1841. Hooker took many specimens of marine fauna from the area, including 22 species of crustacea and molluscs. In addition to describing the specimens taken by the expedition Hooker tried, with some groups of organisms, to compile a list of all the species known to occur in the New Zealand area. Some of the results appeared in 1844-46, but then publication was interrupted by lack of funds, and it was not resumed until 1874.</p>
<p TEIform="p">While Ross and Hooker were exploring the Antarctic seas to the south of New Zealand, they made several other contributions to marine biology. Ross took dredgings from depths as great as 400 fathoms, and by finding living organisms in these dredge-loads he disproved the prevailing theory that life could not exist at such depths. Also, Hooker pointed out the important role of diatoms in the Antarctic Ocean. The biological results from the expedition might have been even more substantial than they were, but, unfortunately, Ross kept all the specimens taken by his dredging in a private collection which disappeared after his death, and so they were never properly examined or described.</p>
<p TEIform="p">At approximately the same time as the Treaty of Waitangi was being signed, an American fleet arrived in the Bay of Islands. This was the first scientific expedition ever sent out by the United States government, and it was accompanied by seven scientists including mineralogist-naturalist James Dana. During his stay in New Zealand, Dana concentrated on the crustacea of the area and found a large number of species including more than 30 which were previously unknown. He also described a few coelenterates from the area. All of his results were published by 1855 in the official report of the expedition.</p>
<p TEIform="p">Huxley spent a week in the Bay of Islands in May 1850 while <hi rend="i" TEIform="hi">H.M.S. Rattlesnake</hi>, on which he was serving as assistant surgeon,
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underwent repairs. He seems to have done little of note, himself, while in New Zealand, but the ship's official naturalist, John Macgillivray, collected a few polyzoans and sertularians which were described by George Busk in an appendix to the official narrative.</p>
<p TEIform="p">During the years 1847 to 1849, <hi rend="i" TEIform="hi">H.M.S. Acheron</hi> was engaged in surveying the coasts of the North and South Islands. The surgeon of the vessel, David Lyall, made a collection of the polyzoa which was sent to the British Museum where it was described by Busk. In 1849 Frederick Strange joined the <hi rend="i" TEIform="hi">Acheron</hi>, and he apparently made several offshore dredgings, the first ever in New Zealand waters (Suter, 1913). Several molluscs were taken in these dredgings, and they were described to the Royal Zoological Society by A. Adams and C. P. Deshayes in the early 1850's.</p>
<p TEIform="p">The last noteworthy expedition of this era was that of the Austrian ship, <hi rend="i" TEIform="hi">Novara</hi>, which visited Auckland and the <name type="person" key="name-110561" TEIform="name">Hauraki</name> Gulf in 1858-59. The <hi rend="i" TEIform="hi">Novara's</hi> naturalists collected some molluscs and other organisms from the area. The descriptions of these were published in Vienna from 1864 to 1869, but care was not taken to separate the New Zealand collections from those from other parts of the voyage, and so confusion ensued.</p>
<p TEIform="p">While the work of these early expeditions may not seem very impressive by modern standards, it provided a solid base on which to construct future research. These exploring naturalists discovered and described a large number of new marine organisms, and, perhaps even more importantly, they stimulated much interest among European scientists in this region of the ‘underside’ of the globe.</p>
</div2>
<div2 id="t1-body-d1-d2" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Early Resident Activity</hi></head>
<p TEIform="p">Even while the early explorers were still sailing through New Zealand waters, a new era in the scientific investigation of the region was unfolding. Beginning in the 1830's, resident scientists, or at least residents with some interest in science, began to appear in New Zealand. This meant that collections and observations could be made over long periods of time as opposed to the investigations of the earlier expeditions which had always been limited to a few weeks or at most a few months. Moreover, these residents were concentrating their efforts almost entirely on New Zealand flora and fauna, while the early explorers saw the islands as being only a small part of the total area of their investigations.</p>
<p TEIform="p">The first European resident to provide information on marine biology was <name type="person" key="name-208989" TEIform="name">J. S. Polack</name>, who lived in New Zealand from 1831 to 1837. In describing his ‘travels and adventures’ in the region, Polack devoted a whole chapter to natural history (Polack, 1838, chap. IX). He wrote in a popular style and did not give any systematic descriptions, but he was a careful observer and made many valuable comments on the ecology and behaviour of animals. In the marine
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field he noted everything from ‘leviathans’ (whales) to ‘large glutinous masses’ of plankton. He also devoted an appendix in his journal to a study of the whales of the area which must rank as one of the best works of that period on their behaviour and ecology (Polack, 1838, vol. II, note 8).</p>
<p TEIform="p">The first professional scientist, other than a ship's naturalist, to visit New Zealand was Dr. Ernst Dieffenbach. In 1838 he was commissioned by the New Zealand Company, and instructed that ‘General information relating to navigation, geography, geology, botany, and zoology, and the traditions, customs, and character of the natives, will be highly appreciated, and will be communicated from time to time to the scientific societies in England’ (Callaghan, 1957). Dieffenbach had all of his expenses paid by the company plus £50 for an outfit and £50 for a premium, and he was promised ‘to be remunerated hereafter by the company according to his exertions’ (Callaghan, 1957). He remained in New Zealand until October 1841, and on his return to England he published an account of his findings which included a section on the fauna of the region.</p>
<p TEIform="p">In addition to bringing home many specimens, Dieffenbach attempted with the help of several British experts to put together a faunal list of all the species known to exist in New Zealand. Based on his work, Sir John Richardson compiled a list of 92 species of New Zealand fish including several new ones found by Dieffenbach. Similarly, J. E. Gray, the keeper of the zoological collections at the British Museum, worked on other groups of New Zealand marine organisms. He came up with 225 species of molluscs, 29 crustacea, and numerous species from other invertebrate groups. These lists and descriptions were published in Dieffenbach's <hi rend="i" TEIform="hi">Travels in New Zealand</hi> in 1843.</p>
<p TEIform="p">Many other men living in New Zealand during this period were making contributions to the knowledge of local marine biology. For example, in 1835 Rev. <name type="person" key="name-209706" TEIform="name">W. Yate</name> sent a collection of mollusc shells, which included 10 new species, to the British Museum. In 1834 Rev. <name type="person" key="name-207684" TEIform="name">William Colenso</name> arrived in the Bay of Islands as a missionary printer. He became one of the outstanding figures among New Zealand botanists, and he also did a considerable amount of work in marine zoology, publishing several papers on molluscs.</p>
<p TEIform="p">Shortly after the middle of the nineteenth century, moves were made to organise and co-ordinate the diverse scientific work being carried out by naturalists in New Zealand. The first attempt to found a scientific organisation in the colony was made by Sir George Grey in 1851, resulting in the establishment of the New Zealand Society. That body was very short lived, however, and it does not appear to have left any substantial legacy. In 1862 Julius Haast set up the Philosophical Institute of Canterbury. Then, in 1867, some of the former members of the New Zealand Society founded the Wellington Philosophical Society. This was followed shortly in 1868 by the
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establishment of the Auckland Philosophical Society. These groups provided local forums in which the scientists of the respective provinces could exchange information and ideas.</p>
<p TEIform="p">At this time the need for a national scientific body was realised, and in 1867 <name type="person" key="name-208190" TEIform="name">James Hector</name> framed the ‘New Zealand Institute Act’ which called for the establishment of ‘an institute for the advancement of Science and Art in New Zealand’. The act was duly passed by the government, and the institute held its first meeting on August 4, 1868. The early members decided that their primary object should be ‘to promote the general study and cultivation of the various branches and departments of art, science, literature, and philosophy’ (Bowen, 1868), and the government agreed to an annual grant of £500 to aid in this pursuit.</p>
<p TEIform="p">The New Zealand Institute (which later became the Royal Society of New Zealand when it received the sponsorship of King George V in 1933) met regularly and published an annual volume of its proceedings and transactions which also contained the proceedings of the provincial societies. This provided scientists from all over the country with a system of communication for their ideas. It was especially crucial for the scientific community in New Zealand to have such a body because of the isolation of the country from the main European centres. As well as providing a vehicle for the exchange of ideas, the institute could also serve to co-ordinate the activities of New Zealand scientists. The important influence of the institute in its early days was pointed out by F. R. Callanghan, a more recent president of the Royal Society, when he said, ‘From 1867 onwards there was a very real trend away from scientific work done in isolation towards that done in association’ (Callaghan, 1952).</p>
<p TEIform="p">While the aim of the institute was to promote science in general, all of the individual disciplines naturally benefited from its efforts, and marine biology was no exception. The first volume of the <hi rend="i" TEIform="hi">Transactions and Proceedings</hi> contained several papers on marine science, and that publication was to continue to be the main forum for the work of New Zealand marine biologists for many years. With the exception of a few papers sent to European societies, virtually all of the research carried out by local marine biologists for the next several decades was communicated through the institute (Freed, 1963).</p>
<p TEIform="p">Closely related to the founding of the scientific societies were the beginnings of another type of establishment that was to prove of value to the marine biologist — the natural history museum. The first of these, the Canterbury Museum, was founded by Haast in 1861, largely made up from his own collections, and it first exhibited to the public in 1865. The Colonial Museum in Wellington (later the ‘Dominion’ and now the ‘National’ Museum) was set up in 1865 mainly through the efforts of Hector. It took over the collections of the defunct New Zealand Society plus some of the specimens from the New Zealand Exhibition held in Dunedin in 1865. The Exhibition
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also led to the founding of the Otago Museum in 1868 to house the remaining Exhibition collections as well as the collections from the provincial geological survey. The Auckland Philosophical Society started a museum in that city in 1867, and in the years following several smaller museums were established in other cities of both the North and South Islands.</p>
<p TEIform="p">These museums soon built up large collections which included many marine specimens, and thereby provided the basic material needed for systematic studies of marine biology. Prior to the establishment of the local museums, the most accessible collections for New Zealanders were in the British Museum. Consequently, most specimens had to be shipped to London for identification at great expense, both in terms of time and money. Thus, the appearance of significant collections in New Zealand clearly was a great stimulus to the development of local marine biology.</p>
<p TEIform="p">Another organisation started during this period which had an influence on marine biology was the Geological Survey. This body, founded in 1865, superseded the earlier provincial societies. It represented the first official attempt by the national government to stimulate scientific effort. It was very closely tied to the Colonial Museum and the New Zealand Institute, partly because the museum was the chief repository for the collections of the Survey, and also because the first director of the Survey, <name type="person" key="name-208190" TEIform="name">James Hector</name>, served as the director of the museum and the general manager of the institute at the same time. While the primary interest of the Survey was geological, it also made many important contributions to marine biology. Several scientists carried out significant marine research while employed by the Survey, the most notable of these being Hector himself and one of his early assistants, <name type="person" key="name-208309" TEIform="name">F. W. Hutton</name>. Furthermore, the Geological Survey supervised the publication of many important biological papers.</p>
<p TEIform="p">The late 1860's saw one more development which had an important effect on the advancement of New Zealand marine biology: the founding of the first university in the country. In June of 1869 the Otago Provincial Council passed the ‘University of Otago Ordinance’, thereby providing New Zealanders with their first opportunity to obtain higher education without going abroad. From a scientific viewpoint the founding of the University of Otago was especially significant because it was one of the first universities in the British Empire to emphasise the teaching of natural science. At the first meeting of the university council in November 1869, <name type="person" key="name-134232" TEIform="name">James Macandrew</name>, the provincial superintendent, advised, ‘While due provision should be made for classical and metaphysical studies, there should be equal, if not greater prominence given to the teaching of Natural Sciences’ (Jenkinson, 1940). His words were heeded, and one of the four founding chairs was for natural science. While at first there was much emphasis on the agricultural and geological sciences, marine biology also had an important place in the early curriculum. It is
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worth noting that the first B.Sc. degree granted in New Zealand was given to a marine biologist, <name type="person" key="name-207643" TEIform="name">Charles Chilton</name>, by Otago in 1887.</p>
<p TEIform="p">In 1870 Parliament passed a ‘New Zealand University Act’ in order to set up a national university system. Canterbury College was the first to appear under the act, being founded in 1873. It was followed by Auckland University College in 1883 and Victoria University College in 1899. With the establishment of this university system, the country was now capable of training its own scientists, and therefore New Zealand marine biology was no longer totally dependent on imported talent.</p>
<p TEIform="p">Within the framework set up by the institutions founded in the 1860's, research began to expand in the early 1870's. Certainly the most prolific and probably the most important worker in marine biology during this period was <name type="person" key="name-208309" TEIform="name">Frederick Wollaston Hutton</name>. He arrived in Auckland from England in 1866 intending to be a farmer, but in 1871 he became an assistant in the Geological Survey in Wellington. Two years later he moved to Otago to become provincial geologist, lecturer at the university, and curator of the Otago Museum. In spite of his vocational orientation toward geology, Hutton did a large amount of work in marine biology, publishing 56 papers on that subject from 1872 to 1896.</p>
<p TEIform="p">Hutton's greatest contributions to the science were his faunal catalogues that appeared in the early 1870's. Based on his own collections as well as all of the previous work done on New Zealand, he compiled descriptive lists of all the known animals in certain groups. In 1872 he published, under the auspices of the Colonial Museum and the Geographical Survey, <hi rend="i" TEIform="hi">The Fishes of New Zealand</hi> in which he described the 141 species of fish known at that time to exist in New Zealand waters. This was followed by the <hi rend="i" TEIform="hi">Catalogue of Echinoderms in New Zealand</hi>, also published in 1872. Hutton's expertise as a naturalist is shown by the fact that of the 34 species of echinoderms in the catalogue only 10 had previously been described from New Zealand waters. Hutton himself found and described 18 new species and six others never before recorded in New Zealand. In 1873 he published the <hi rend="i" TEIform="hi">Catalogue of the Marine Mollusca of New Zealand</hi> which included descriptions of more than 300 species of molluscs, plus numerous other organisms such as polyzoa, brachiopods, and tunicates which are now classified in other phyla. These catalogues were later incorporated into Hutton's magnum opus, the <hi rend="i" TEIform="hi">Index Faunae Novae Zelandiae</hi>, published in 1904.</p>
<p TEIform="p">Hutton did not stand alone at this time. His fellow geologist, Hector, also published more than 50 papers between 1869 and 1901 on subjects related to marine biology (Freed, 1963), and many others were active in the field. The rapid advance of the science during the period was shown by <name type="person" key="name-208417" TEIform="name">T. W. Kirk</name> when he pointed out that from 1850
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to 1875 the number of species of fish known to inhabit New Zealand waters increased from 92 to 163, molluscs from 240 to 502, and whales from four to 21 (Kirk, 1875).</p>
<p TEIform="p">No description of the history of marine biology in any part of the globe would be complete without some mention of the voyage of <hi rend="i" TEIform="hi">H.M.S. Challenger.</hi> Many people today consider the <hi rend="i" TEIform="hi">Challenger</hi> expedition to be the beginning of ‘modern’ oceanography. The results obtained by this British ‘spar-decked corvette’ certainly represent a major milestone in marine biology. Under the command of C. Wyville Thomson, the <hi rend="i" TEIform="hi">Challenger</hi> circumnavigated the globe along a track of nearly 69,000 miles from December 1872 to May 1876. Soundings and dredgings were made at 362 different stations, and specimens were taken from depths down to 4,475 fathoms. The results of the voyage occupied the attention of experts from all over the world for 19 years after the ship's return, and when finally publishd they filled 50 quarto-sized volumes. Thirty-four of these volumes were devoted entirely to marine biology. By modern reckoning the <hi rend="i" TEIform="hi">Challenger</hi> brought back 4,417 species of animals which had never before been seen. Her findings provided ‘an absolute beginning’ for several areas of marine biology and caused a ‘complete revision’ of many existing fields of study (Lankester <hi rend="i" TEIform="hi">et al.</hi>, 1895).</p>
<p TEIform="p">The <hi rend="i" TEIform="hi">Challenger</hi> visited New Zealand in June and July 1874, and, as in every area visited, she made a substantial contribution to the knowledge of the local seas. The <hi rend="i" TEIform="hi">Challenger</hi> crew worked seven stations in New Zealand waters, and dredgings from more than 100 fathoms were made at all of them. This work represented the first serious investigation of the deep-sea fauna of the area. When the results of these dredgings were examined, more than 220 new species were found. The descriptions of these organisms were published both in the official reports of the expedition and in special articles published in New Zealand (Watson, 1883; Hamilton, 1896).</p>
<p TEIform="p">At the same time as the <hi rend="i" TEIform="hi">Challenger</hi> was off New Zealand, a French expedition was observing a transit of Venus from the Campbell Islands. The naturalist with the expedition, H. Filhol, made some dredgings from 20 to 35 fathoms in various areas including Cook Strait, Foveaux Strait, and off the Auckland and Campbell Islands (Yaldwyn, 1957). Filhol later toured parts of the North and South Islands making further collections. Between 1878 and 1886 he published several papers on the invertebrate fauna of the areas he visited, the most well-known of which was a catalogue of the New Zealand crustacea.</p>
<p TEIform="p">Another European scholar examining New Zealand crustacea during this period was E. J. Miers of the British Museum. He began his work by looking at the crustacea from the <hi rend="i" TEIform="hi">Erebus</hi> and <hi rend="i" TEIform="hi">Terror</hi> expedition but then extended his search for New Zealand specimens to the whole of the British Museum collections. In all, Miers found and described 140 species of New Zealand crustacea, and he published
<pb id="n13" n="200" TEIform="pb"/>
this work in 1876. This research proved so valuable and so durable that Chief Fisheries Inspector, A. E. Hefford, writing in 1947, was able to say, ‘No comprehensive monograph on crustacea has yet been written to succeed Miers’' (Hefford, 1947).</p>
<p TEIform="p">The 1880's and 1890's lacked any spectacular advances in marine biology, but throughout that period New Zealanders were doing a considerable amount of sound research in the discipline. <name type="person" key="name-208922" TEIform="name">T. J. Parker</name>, who in 1880 was appointed Professor of Biology at Otago University and curator of the museum, carried out numerous investigations into New Zealand fishes. <name type="person" key="name-207639" TEIform="name">T. F. Cheeseman</name> was exploring the marine fauna around Auckland during this same period. Although primarily a botanist, he did some dredging and published several papers on the molluscs of that area. Another versatile scientist, <name type="person" key="name-208417" TEIform="name">T. W. Kirk</name>, was examining marine life during these decades. He started his scientific career as a cadet with the Geological Survey, and he finished it as the head of the Horticulture Division of the Department of Agriculture, but between 1879 and 1888 he managed to produce more than 10 papers on several different aspects of marine zoology. Arthur Dendy, who learned much of his biology from working on the specimens brought back by the <hi rend="i" TEIform="hi">Challenger</hi>, came to Canterbury as a lecturer, and later professor, in zoology. Before he left to go to South Africa in 1905, he contributed a great deal to the knowledge of New Zealand sponges. In addition to these men, many others were working in the field during this period, and the volumes of the <hi rend="i" TEIform="hi">Transactions and Proceedings of the New Zealand Institute</hi> for these years contain numerous reports of their findings in marine biology.</p>
</div2>
<div2 id="t1-body-d1-d3" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">From the Turn of the Century</hi></head>
<p TEIform="p">Around the turn of the century, several events took place which held great importance for the development of marine biology in New Zealand. The most important of these was the establishment of the country's first laboratory for marine research at Portobello, near Dunedin.</p>
<p TEIform="p">The need for a marine station was first noted by <name type="person" key="name-208922" TEIform="name">T. J. Parker</name> in his presidential address to the Otago Institute in 1886 (Parker, 1886). He called on the government to establish a laboratory ‘for the purpose of developing our fisheries’. The cause was again taken up by <name type="person" key="name-209458" TEIform="name">G. M. Thomson</name> in another address to the Otago Institute in October 1895 in which he expounded the potential benefits of such a laboratory, especially for the introduction of foreign food fishes to New Zealand. The institute was ‘sufficiently impressed’ to form a committee to investigate the project and to search for a suitable site in the summer of 1895-96. The issue was placed before the government, and in 1897 Parliament voted a grant of £750 ‘For fish-hatcheries and expenses of Expert Ayson to Canada and America’ (Thomson, 1921). While the government's ‘expert’, <name type="person" key="name-207310" TEIform="name">L. F. Ayson</name>, was examining fish hatcheries in
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North America and Europe, the Otago Institute, aided by the Otago Acclimatisation Society, drew up plans and started to raise money for the laboratory. A governing board was established, made up of members from Otago University, the institute, the acclimatisation society, and the government, which was to report annually to the Minister of the Marine.</p>
<p TEIform="p">There followed a period of ‘negotiation with the government and protracted delay’ (Thomson, 1921). The governing board finally held its first official meeting in June 1902, and Thomson was elected president. Construction of the laboratory on the chosen site at Portobello began in early 1903, and the buildings were formally opened on January 13, 1904. The station consisted of an aquarium and laboratory block, a four-room cottage, a small pump house, and two large fish ponds. The total cost of construction was £1,448, of which £800 came from the government, £250 each from the Otago Institute and the acclimatisation society, and the rest from other organisations and individuals from around New Zealand.</p>
<p TEIform="p">The early work of the marine station was largely devoted to the introduction of European food fishes into New Zealand seas. Attempts were made to introduce turbot, crabs, and lobsters, but these were not successful. About the time of the First World War, those in charge of the laboratory began to realise the futility of their acclimatisation attempts, and they shifted the emphasis of their research. Several projects were then initiated which were to have lasting value. Among these were crustacean studies by Thomson, temperature recordings and drift bottle experiments off the Otago coast, faunal surveys of Otago Harbour, and much general taxonomic work. A fish tagging project on sole was attempted in 1916, but it was thwarted by the lack of a suitable material for the tags (Thomson, 1921).</p>
<p TEIform="p">During this period the government gave an annual grant of £200 to provide for a research assistant at the marine station. In 1921 M. W. Young was appointed as the first resident marine biologist (previously the station had been manned only by a curator). When Young moved to Wellington in 1926. <name type="person" key="name-031343" TEIform="name">D. H. Graham</name> replaced him. Significant research continued at the laboratory until the early 1930's when the general economic depression caused the government to reduce its financial support. Consequently, in 1932 the biologist and his assistant had to be dismissed because of lack of funds. To make matters worse, <name type="person" key="name-209458" TEIform="name">G. M. Thomson</name>, who had been the guiding spirit of the laboratory since its inception, died in 1933. To quote the recent director of the station, Dr. <name type="person" key="name-207364" TEIform="name">E. J. Batham</name>, ‘The story from then until 1951 was one of steady deterioration’ (Batham, 1956).</p>
<p TEIform="p">Another significant occurrence around the turn of the century was the birth of local interest in the offshore and deep-water fauna of the region. In 1893 Capt. J. Fairchild of <hi rend="i" TEIform="hi">N.Z. G. S. Stella</hi> undertook some dredging in 110-170 fathoms off the Bounty Islands. Very little is known about the results of this work, and nothing was published
<pb id="n15" n="202" TEIform="pb"/>
until 1933 when <name type="person" key="name-209010" TEIform="name">A. W. B. Powell</name> described some of the molluscs taken by Fairchild's dredge (Yaldwyn, 1957). The first locally organised scientific trawling expedition took place in 1899 when ‘a party of gentlemen’ led by <name type="person" key="name-207408" TEIform="name">W. B. Benham</name> set out in the Otago Harbour Board tug <hi rend="i" TEIform="hi">S.S. Plucky.</hi> They trawled in 10 to 30 fathoms off the Otago coast, and they captured a few species of molluscs, crustacea, echinoderms, and annelids (Benham, 1899).</p>
<p TEIform="p">In 1904, on the occasion of its meeting in Dunedin, the Australasian Association for the Advancement of Science appointed a committee ‘to initiate a biological and hydrographical survey of the continental shelf of New Zealand by dredging and sounding’ (A.A.A.S., 1905). It consisted of six prominent marine scientists with <name type="person" key="name-207643" TEIform="name">Charles Chilton</name> as secretary. A grant of £50 was placed at their disposal. The committee wasted little time in getting to work, and they made their first dredging attempt off Port Chalmers on January 14, 1904. The weather was not co-operative, however, and the scientists found themselves ‘in great discomfort’. The inclement weather thwarted the dredging, and a member of the party reported, ‘As a result we lost considerable gear and obtained only a cupful of bottom from 100 fathoms’ (Hedley, 1905).</p>
<p TEIform="p">The A.A.A.S. committee tried again a week later, this time out of Auckland to the east of Great Barrier Island. The weather was not much better, but they did manage to make two dredgings in 110 fathoms. From these hauls they obtained one-third of a ton of ‘soft, sticky, green mud’ in which they found a few echinoderms and molluscs (Hedley, 1905). After these early attempts, the committee was unable to get together to do any further work as a body, but Chilton continued to report to the A.A.A.S. on marine research in New Zealand until the committee was officially disbanded in 1928.</p>
<p TEIform="p">The Philosophical Institute of Canterbury organised an expedition to the Auckland and Campbell Islands in November 1907 on the government steamer <hi rend="i" TEIform="hi">Hinemoa.</hi> They made observations of the littoral fauna of the Sub-Antarctic islands and also made dredgings from depths of 8 to 85 fathoms. The most noteworthy results were from the investigations of the foraminifera made by <name type="person" TEIform="name">F. Chapman</name>. He described 168 species of which 103 had not previously been found in New Zealand waters. Collections of other organisms were examined by the appropriate experts from around the country, and many species new to science were recorded (Chilton, 1909).</p>
<p TEIform="p">In 1908 a private party on the steamer <hi rend="i" TEIform="hi">Rakiura</hi> dredged in 100 and 170 fathoms off Puyseger Point in Southland, but the results appear to have been minimal. Only a few mollusc shells and some foraminiferal sand were taken by the dredge (Yaldwyn, 1957). Unfortunately, this expedition marked the end of locally organised dredging operations for quite a few years.</p>
<p TEIform="p">The early part of the twentieth century saw steady progress being made by New Zealanders in almost all areas of marine biological
<pb id="n16" n="203" TEIform="pb"/>
research. From the time of Quoy and Gaimard on the <hi rend="i" TEIform="hi">Astrolobe</hi>, conchology had always been at the forefront of marine science in New Zealand, and this trend continued after 1900. <name type="person" key="name-209371" TEIform="name">Henry Suter</name>, who came to this country from Switzerland in 1887, began to work on local molluscs shortly after the beginning of the century. He compiled descriptions of all the known specimens of molluscs in the region, and in 1913 he published his <hi rend="i" TEIform="hi">Manual of New Zealand Mollusca.</hi> This work contained descriptions of 1,079 species, and it also served to bring the nomenclature used in New Zealand into conformance with international standards. Suter's manual is still viewed as the classic reference source in New Zealand conchology to the present day.</p>
<p TEIform="p">Similar, though less spectacular, advances were being made in the studies of other groups of marine animals at this time. <name type="person" key="name-207643" TEIform="name">Charles Chilton</name>, with help from <name type="person" key="name-209458" TEIform="name">G. M. Thomson</name> and others, was working on crustacea, while <name type="person" key="name-207408" TEIform="name">W. B. Benham</name>, A. Dendy and H. Farquhar were doing research on coelenterates and echinoderms. <name type="person" key="name-208414" TEIform="name">H. B. Kirk</name> followed Dendy into the study of the porifera and did a great deal to advance the knowledge of New Zealand sponges. There were also numerous workers examining the fishes of the area, the most notable of whom were E. R. Waite, L. T. Griffin and W. J. Phillips.</p>
<p TEIform="p">The flora of the New Zealand seas was an almost untouched field until R. M. Laing began his career. Between 1890 and 1940 he published more than 20 papers on the marine algae of the region, thereby opening up the field for further research. The first work on local marine ecology also appeared during this period in <name type="person" key="name-208879" TEIform="name">W. R. B. Oliver</name>'s paper, <hi rend="i" TEIform="hi">Marine Littoral Plant and Animal Communities in New Zealand</hi>, published in 1923.</p>
<p TEIform="p">Despite the advances by local workers, New Zealand marine biology still received much of its impetus from foreign expeditions in the first part of the twentieth century. After the <hi rend="i" TEIform="hi">Challenger</hi> left New Zealand there was a lapse of many years before another European scientific voyage investigated the area. Foreign interest in the region was renewed, however, about 1910, and for a little more than two decades after that a series of expeditions made important contributions to the marine sciences in New Zealand.</p>
<p TEIform="p">The first of these visitors was the British <hi rend="i" TEIform="hi">Terra Nova</hi> expedition of 1910-13. The main purpose of this voyage was the exploration of Antarctica (which led to Scott's fatal journey to the South Pole), but in 1911 the vessel made a winter cruise around the north of New Zealand. During the course of that cruise, 80 plankton samples were taken, and seven dredgings were made from depths of 15 to 300 fathoms. These revealed ‘a bottom fauna of extraordinary variety, including a great number of forms new to science’ (<hi rend="i" TEIform="hi">British Antarctic Expedition</hi>, 1924). Later the expedition took 135 plankton samples and 50 bottom hauls between the southern part of New Zealand and McMurdo Sound. The zoological reports of the expedition were published by the British Museum in 1924-30, filling eight volumes.</p>
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<p TEIform="p">In the summer of 1914-15, Dr. Th. Mortensen came to New Zealand while on his ‘Pacific Expedition 1914-1916’. Accompanied by Chilton and others, he travelled aboard the government steamer <hi rend="i" TEIform="hi">Hinemoa</hi> on its annual cruise to visit the lighthouses of the North Island. ‘By kind permission of the Minister of Marine’, they were allowed to dredge occasionally during the trip (Chilton, 1921). Mortensen also accompanied <hi rend="i" TEIform="hi">G.S. Amokura</hi> to the Auckland and Campbell Islands. When Mortensen and his colleagues in Copenhagen examined his material from New Zealand, they produced many important results. Among these were the first comprehensive account of the echinoids of the region (Mortensen, 1922) and the discovery of 29 new species of molluscs (Ohdner, 1924). A series of other papers on the marine fauna of New Zealand collected by Mortensen appeared in the <hi rend="i" TEIform="hi">Videnskadelige Meddelelsen fra Danske Naturhistorrisk Forering i Kobenhavn</hi> throughout the 1920's.</p>
<p TEIform="p">Another Danish expedition, the Carlsberg Foundation Oceanographic Expedition around the world 1928-30 on the <hi rend="i" TEIform="hi">Dana</hi>, arrived in Auckland in December 1928. From there the <hi rend="i" TEIform="hi">Dana</hi> proceeded to Wellington and thence to the waters to the east of the South Island. She worked 27 stations in the New Zealand region, ranging from the Kermadecs to the latitude of Stewart Island, and dredgings were taken from depths as great as 3,190 metres. The results were published in the 14 volumes of the <hi rend="i" TEIform="hi">Dana Reports</hi> in Copenhagen in the 1930's.</p>
<p TEIform="p">The ‘British, Australia, and New Zealand Antarctic Expedition’ of 1929-31 did a small amount of work in the New Zealand area. Several dredgings were made around Macquarie Island with hauls taken from depths down to 280 metres, and townet and littoral collections were made in the area as well.</p>
<p TEIform="p">British scientists returned to New Zealand in August 1932 on the <hi rend="i" TEIform="hi">R.R.S. Discovery II.</hi> They took new samples from some of the bottom stations examined by the <hi rend="i" TEIform="hi">Terra Nova</hi> and investigated some additional areas suggested by <name type="person" key="name-209010" TEIform="name">A. W. B. Powell</name> of the Auckland Museum. They used a newly designed conical dredge which was more efficient than any equipment previously used in the area. Therefore, even though most of their stations had already been investigated by the <hi rend="i" TEIform="hi">Terra Nova</hi>, the <hi rend="i" TEIform="hi">Discovery</hi> crew brought up much new material, including 6 new genera and 128 new species of molluscs (Powell, 1937). Several bathypelagic hauls with various types of nets at depths down to 550 fathoms were also made, and these provided other interesting results (Yaldwyn, 1957). This work was followed by another hiatus of European exploration of the area, and no more major foreign expeditions ventured into the New Zealand seas until the early 1950's.</p>
</div2>
<div2 id="t1-body-d1-d4" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Fisheries and Other Government Influence</hi></head>
<p TEIform="p">Throughout its history, New Zealand marine biology has been strongly affected by the fisheries. The desire to capture food from the
<pb id="n18" n="205" TEIform="pb"/>
sea has provided steady encouragement to the scientific investigation of the oceans. The government first took an interest in the problem of the fisheries in February 1869 when it appointed a commission headed by Hector to examine the fisheries of New Zealand. The commission, however, was given no financial support, and for this reason it accomplished little. As one contemporary described their results, ‘All that was learned was what fishmongers and fishermen could tell’ (Sherrin, 1886).</p>
<p TEIform="p">In 1877 J. Macandrew, M.P. from Dunedin, sought legislation to provide protection for flounder and sole during their spawning season. As a result of his efforts, the Fish Protection Act of 1877 was passed, providing the first legal regulation of fishing in New Zealand marine waters. The responsibility for enforcing this act was rather vague, but it apparently fell to the Marine Department. As later acts more clearly spelled out this responsibility, the Marine Department gradually took complete charge of regulating the fishing industry. In 1899, after his return from the tour of overseas fish hatcheries and marine stations, <name type="person" key="name-207310" TEIform="name">L. F. Ayson</name> was appointed Inspector of Fisheries.</p>
<p TEIform="p">Prior to 1900 the Marine Department had sponsored very little scientific enquiry into fisheries problems. In 1890 the department arranged for lighthouse keepers to report their observations on fishes to <name type="person" key="name-209458" TEIform="name">G. M. Thomson</name>, but that appears to be its only scientific contribution in that period. When Ayson took office, he realised the need for more comprehensive fisheries research, and in 1900 he organised an experimental trawling expedition. The government chartered the steam trawler <hi rend="i" TEIform="hi">Doto</hi> from March to June 1900 and set about trawling in various areas off the South Island. On this voyage 154 hauls of the net were made from two to 50 fathoms. The results were sufficiently interesting to warrant further investigation, and in 1901 first the <hi rend="i" TEIform="hi">Doto</hi> and then the <hi rend="i" TEIform="hi">Rita</hi> were chartered to trawl around the North Island. More than 120 hauls from depths of three to 38 fathoms were made during these investigations. At different times in the course of these cruises, prominent marine biologists such as Benham, Thomson and A. Hamilton accompanied the vessels, and they examined and described many of the organisms taken (Waite <hi rend="i" TEIform="hi">et al.</hi>, 1909).</p>
<p TEIform="p">As a follow-up to the 1900-01 expedition, the government chartered the <hi rend="i" TEIform="hi">Nora Niven</hi> from the Napier Fish Supply Company in 1907. The vessel made two cruises for the Marine Department between June and December. During that time she covered most of the coastal areas of both the North and South Islands and visited the Chatham Islands as well. A total of 252 hauls were taken from depths down to 105 fathoms. The main purpose of the voyage was commercial fisheries research, but E. R. Waite of the Canterbury Museum was invited on board to represent the scientific community. Unfortunately, because of the fisheries orientation of the expedition, means were not provided to capture small invertebrates in the trawls. Also, although many plankton samples were netted, all of the specimens were washed
<pb id="n19" n="206" TEIform="pb"/>
overboard during a storm. In spite of these problems, many organisms were collected, and the results were analysed by experts including Suter, Benham, Chilton and Waite. Among the invertebrates five new species of echinoderms and eight new molluscs were described. Furthermore, Waite described 85 species of fish, 11 of which were new (Waite <hi rend="i" TEIform="hi">et al.</hi>, 1909). As late as 1947, Waite's report on the <hi rend="i" TEIform="hi">Nora Niven</hi> expedition was still considered to ‘represent the most complete and comprehensive monograph on New Zealand ichthyology yet published’ (Hefford, 1947).</p>
<p TEIform="p">In a report in 1913, Ayson pressed for more effort on scientific fisheries research, but apparently his advice was ignored. The following year Prof. E. P. Prince from the Commission of Fisheries for Canada was brought to New Zealand to report on the fisheries. After spending four months examining local fishing, he agreed with Ayson that more research was needed. Prince specifically recommended that a marine laboratory-hatchery comparable to Portobello be established on the North Island and that greater use be made of the four universities for fish research. This advice also appears to have gone unheeded for many years by the government (Martin, 1969).</p>
<p TEIform="p">The scientific capabilities of the Marine Department were improved in 1925 when A. E. Hefford joined the staff as a fisheries expert. He had worked in several biological institutions in Britain and was well versed in the problems of fisheries conservation. Hefford became Chief Inspector of Fisheries in 1927, and during the 20 years he held that post, he did much to encourage marine research. He also had an influence through other scientific organisations, especially the Wellington Philosophical Society which he served as president from 1937 to 1939.</p>
<p TEIform="p">In 1926 the Fisheries Branch of the Marine Department acquired the services of its first full-time marine biologist, M. W. Young, from Portobello. As Hefford points out, ‘The time had arrived when it had to be recognised that a basis of biological fact was essential for the proper understanding of the problems of fisheries administration’ (Hefford, 1947). Young did a considerable amount of work on oysters, especially those of the Auckland region. In 1936 Young was given an administrative post, and he was succeeded as marine biologist by A. M. Rapson. Rapson worked on the distribution and life history of several commercially important fishes. In 1938 he accompanied the <hi rend="i" TEIform="hi">Discovery II</hi> on a voyage to Antarctica and gained valuable training in modern methods of biological investigation.</p>
<p TEIform="p">Despite the appointment of a marine biologist, the Fisheries Branch was still a long way from being an efficient research unit at that time. In his official report for 1929 Hefford was forced to complain,</p>
<p TEIform="p">In the absence of a research staff suitably equipped with laboratory accommodation and with facilities for pursuing
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investigations at sea, the rudimentary branch of our organisation concerned with scientific research cannot be expected to produce any monumental results. (Hefford, 1929)</p>
<p TEIform="p">In 1937 they were provided with a laboratory on Sydney Street in Wellington, but even that proved to be less than ideal. As for field work, most fisheries researchers were limited to accompanying commercial fishermen on their regular voyages. After much effort the government was convinced in the late 1930's to provide a fisheries research vessel, but the timing proved to be very poor. The 65-foot research ship <hi rend="i" TEIform="hi">Ikatere</hi> was launched just in time to be commandeered for an auxiliary vessel in the Second World War. Finally, when the war was over, the ship was refitted for fisheries research.</p>
<p TEIform="p">Government sponsorship of marine research was also to come through channels other than the Fisheries Branch of the Marine Department. In 1926 Sir Frank Heath, the founder of the British Department of Scientific and Industrial Research, visited New Zealand and wrote a report on the state of science in the country. Based on his recommendations, an act was passed in October 1926 ‘to make provision for the promotion and organisation of scientific research, and for its application to the primary and secondary industries of New Zealand’. This act led to the establishment of the New Zealand Department of Scientific and Industrial Research which brought together under one organisation most government sponsored research activities. As will be seen below, the D.S.I.R. came to have a strong influence on the development of the marine sciences in New Zealand.</p>
</div2>
<div2 id="t1-body-d1-d5" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Post-War Development</hi></head>
<p TEIform="p">The end of the Second World War found New Zealand marine biology in a somewhat sorry condition. Fisheries work had fallen off greatly during the war, and by 1947 there was still only one marine biologist on the payroll of the Marine Department. The only institution in the country at this time which was entirely concerned with marine biological research was the laboratory at Portobello, and it was in ‘a state of almost suspended animation’ (Hefford, 1947). Fortunately, this state of affairs did not last long.</p>
<p TEIform="p">During the war, observation and communication parties had been stationed on the Auckland and Campbell Islands as a defence precaution These parties made many observations which were to prove of value to marine biologists, including daily hydrological records from 1941 to 1946 and collections of the fauna of the area. Their results occupied marine biologists for almost a decade and led to a series of publications under the auspices of the D.S.I.R. and the Dominion Museum.</p>
<p TEIform="p">In 1946 the D.S.I.R. sponsored an expedition to the southern fiords on the <hi rend="i" TEIform="hi">New Golden Hind.</hi> During the expedition 84 bottom samples were taken with a Peterson grab from depths of two to 73 fathoms.</p>
<pb id="n21" n="208" TEIform="pb"/>
<p TEIform="p">Much material of interest was found in these samples, including 342 species of molluscs, 17 of which were new (Fleming, 1950).</p>
<p TEIform="p">Before 1950, New Zealand marine sciences suffered from a dearth of organisation and co-ordination. An attempt to organise the discipline had been made in 1927 with the formation of the ‘committee on the oceanography of the Pacific for New Zealand’, but it proved unsuccessful. This lack of organisation became apparent following the arrival of the surveying ship <hi rend="i" TEIform="hi">H.M.S. Lachlan</hi> in 1949. When her commanding officer, Commander Sharpley-Schafer, offered to undertake collections of oceanographic materials, several institutions expressed interest, but they soon discovered that there was no body which could co-ordinate their various needs and desires. Consequently, an ‘Interdepartmental Committee on Oceanography’ was formed in October 1949 with representatives from Victoria University College, the Dominion Museum, the Marine Department, and the D.S.I.R. The committee provided equipment for the <hi rend="i" TEIform="hi">Lachlan</hi> and arranged for the examination of all the specimens taken by the vessel.</p>
<p TEIform="p">In April 1950 the Interdepartmental Committee resolved to encourage the formation of a more comprehensive national body in order to further the co-ordination efforts which they had started. A ‘New Zealand Oceanographic Committee’ was established under the control of the D.S.I.R. in October 1950. It consisted of representatives from the four universities, the Auckland Institute and Museum, the Dominion Museum, the Portobello Marine Biological State, the Meteorological Office, the Marine Department, and the Navy. This new committee was designed to ‘co-ordinate, correlate, and assist oceanographic work in New Zealand’ (Knox, 1953).</p>
<p TEIform="p">By the early 1950's, marine biology in New Zealand had a solid foundation from which to carry out future research. A survey of that period (Knox, 1953) found ‘over 20’ workers in the field of marine taxonomy, and comparable numbers were attacking other areas of marine research. Similarly, many new expeditions, of both foreign and domestic origin, began to comb the New Zealand seas at this time. Their accomplishments are too numerous to be described here. All that can really be done within the scope of this paper is to indicate some of the trends and organisations which have recently furthered the development of the science.</p>
<p TEIform="p">In 1949 a Geophysical Observatory was established in the D.S.I.R. to carry out studies of the physical properties of the ocean. It became part of the Geophysics Division of the D.S.I.R. in 1951 and was renamed the Oceanographic Observatory. Proposals emanating from the Commonwealth Conference on Oceanography in 1954 led to the creation of the New Zealand Oceanographic Institute as a branch of the Geophysics Division. This new institute consolidated all of the government oceanographic research bodies. In 1958 the Oceanographic Institute became a separate branch of the D.S.I.R. Throughout its
<pb id="n22" n="209" TEIform="pb"/>
brief history, the institute has sponsored important research into marine biology as well as into other aspects of New Zealand oceanography (D.S.I.R., 1960).</p>
<p TEIform="p">Laboratory facilities have also improved greatly since 1950. The Portobello marine station was taken over by Otago University in 1951. Under the direction of Dr. <name type="person" key="name-207364" TEIform="name">E. J. Batham</name>, it has been greatly rehabilitated, and much of the facility has been rebuilt and improved. The early ‘sixties saw the opening of three new marine laboratories: Canterbury University's Edward Percival Laboratory at Kaikoura, Auckland University's marine station at Leigh, and Victoria University's marine laboratory at Island Bay. These not only provided great opportunities for student work, but they also enabled research to be carried out in different sections of the country.</p>
<p TEIform="p">Several new organisations and publications were founded during this period which helped to widen communication among New Zealand biologists. In 1952 the New Zealand Ecological Society was established, and much of its attention has been focussed on the problems of marine ecology. In 1960 the founding of the New Zealand Marine Sciences Society provided the first national society entirely dedicated to the advancement of marine research. Its annual meetings provide an important forum for the exchange of ideas among the nation's marine scientists. Most recently, the creation of the <hi rend="i" TEIform="hi">New Zealand Journal of Marine and Freshwater Research</hi> in 1967 has added yet another channel of communication for marine biologists.</p>
<p TEIform="p">Fisheries research has also undergone significant changes in recent years. In December 1964, on the recommendation of the National Research Advisory Council, all fisheries research was consolidated in a new division of the Marine Department known as the Fisheries Research Division. Then, in 1972, control of this division was shifted to the Ministry of Agriculture. By that time the division employed a graduate staff of more than 20 and had greatly expanded facilities including a new laboratory in Christchurch.</p>
<p TEIform="p">All of these institutions provide New Zealand marine biologists with the tools they need to carry out their trade. Furthermore, there has recently been considerable interest in marine biology among students at local universities, and so new researchers are constantly entering the field. This improvement of facilities and increase in interest certainly bodes well for the future of the science in New Zealand.</p>
<p TEIform="p">The changes wrought to the study of marine biology in New Zealand during the slightly more than 200 years since the arrival of Capt. Cook have been remarkable. The first European explorers brought the techniques of the naturalist to an area which had previously been ruled by primitive superstitions. Early residents then continued the study of the marine environment, and soon established the organisational framework necessary for the expansion of the science. Within this framework New Zealand scientists were able to
<pb id="n23" n="210" TEIform="pb"/>
accomplish a tremendous amount, but they also received important inputs from foreign scientific expeditions visiting the area. Toward the end of the nineteenth century, the government began to support and encourage the study of marine biology. It is this structure made up of an active local scientific community, supported by the government, and receptive to the contributions of foreign scientists, that has advanced New Zealand marine biology to the high level where it stands today.</p>
</div2>
<div2 id="t1-body-d1-d6" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Acknoweldgements</hi></head>
<p TEIform="p">I wish to thank the staff of the Zoology Department at Victoria University of Wellington whose kindness and hospitality greatly encouraged me in this project. Foremost among them in deserving thanks is Professor <name type="person" key="name-170438" TEIform="name">J. A. F. Garrick</name> without whose help I could never have written this paper. I would also like to express my gratitude to Dr. R. <name type="person" TEIform="name">J. McIntyre</name> for his comments and criticisms on an early draft of the paper.</p>
</div2>
<div2 id="t1-body-d1-d7" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Bibliography</hi></head>
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<bibl default="NO" TEIform="bibl">Freed, D., 1963: <hi rend="i" TEIform="hi">Bibliography of New Zealand Marine Zoology 1769-1899.</hi> New Zealand Department of Scientific and Industrial Research Bulletin 148. New Zealand Oceanographic Institute Memoir 16.</bibl>
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<bibl default="NO" TEIform="bibl"><name type="person" key="name-208241" reg="T. M. Hocken" TEIform="name">Hocken, T. M.</name>, 1896: Abel Tasman and his Journal. <hi rend="i" TEIform="hi">Trans. Proc. N.Z. Inst.</hi> 28: 117-140.</bibl>
<bibl default="NO" TEIform="bibl"><seg id="s211_3" part="N" TEIform="seg"><name type="person" key="name-208309" TEIform="name">Hutton, F. W.</name></seg>, 1872: <hi rend="i" TEIform="hi">Catalogue of the Echinodermata of New Zealand.</hi> Colonial Museum and Geological Survey Department, Wellington.</bibl>
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<bibl default="NO" TEIform="bibl"><seg sameAs="s211_3" part="N" TEIform="seg">——,</seg> 1873: <hi rend="i" TEIform="hi">Catalogue of the Marine Mollusca of New Zealand.</hi> Colonial Museum and Geological Survey Department, Wellington.</bibl>
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<bibl default="NO" TEIform="bibl">Jenkinson, S. H., 1940: <hi rend="i" TEIform="hi">New Zealanders and Science.</hi> Department of Internal Affairs, Wellington.</bibl>
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<bibl default="NO" TEIform="bibl"><name type="person" key="name-170427" reg="G. A. Knox" TEIform="name">Knox, G. A.</name>, 1953: Oceanographic Research in New Zealand. <hi rend="i" TEIform="hi">7th Sci. Cong. Rep. R. Soc. N.Z.</hi> 72-80.</bibl>
<bibl default="NO" TEIform="bibl">Lankester, E. R., <hi rend="i" TEIform="hi">et al</hi>, 1895: The Scientific Results of the ‘Challenger’ Expedition. <hi rend="i" TEIform="hi">Nat. Sci.</hi> 7: 7-75.</bibl>
<bibl default="NO" TEIform="bibl"><seg id="s211_4" part="N" TEIform="seg"><name type="person" key="name-208623" TEIform="name">McNab, R.</name></seg>, 1907: <hi rend="i" TEIform="hi">Murihiku.</hi> Whitcombe and Tombs Ltd., Wellington.</bibl>
<bibl default="NO" TEIform="bibl"><seg sameAs="s211_4" part="N" TEIform="seg">——,</seg> 1914: <hi rend="i" TEIform="hi">From Tasman to Marsden.</hi> J. Wilke and Co., London.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-121463" reg="E. R. Martin" TEIform="name">Martin, E. R.</name>, 1969: <hi rend="i" TEIform="hi">Marine Department Centennial History 1866-1966.</hi> Marine Department, Wellington.</bibl>
<bibl default="NO" TEIform="bibl">Miers, E. J., 1876: <hi rend="i" TEIform="hi">Catalogue of Stalk and Sessile Eyed Crustacea of New Zealand.</hi> British Museum, London.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-208768" reg="W. P. Morrell" TEIform="name">Morrell, W. P.</name>, 1958: <hi rend="i" TEIform="hi">Sir <name type="person" key="name-123818" TEIform="name">Joseph Banks</name> in New Zealand.</hi> A. H. and <name type="person" key="name-209054" TEIform="name">A. W. Reed</name>, Wellington.</bibl>
<bibl default="NO" TEIform="bibl">Mortensen, T., 1922: Echinoderms of New Zealand and the Auckland-Campbell Islands. <hi rend="i" TEIform="hi">Vidensk. Meddr. dansk. naturh. Foren.</hi> 73: 139-198.</bibl>
<bibl default="NO" TEIform="bibl">Murray, J., and Hjort, J., 1912: <hi rend="i" TEIform="hi">The Depths of the Ocean.</hi> MacMillan and Co. Ltd., London.</bibl>
<bibl default="NO" TEIform="bibl">Odhner, N. H., 1924: New Zealand Mollusca. <hi rend="i" TEIform="hi">Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16</hi>, Vol. 19. Copenhagen.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-208879" reg="W. R. B. Oliver" TEIform="name">Oliver, W. R. B.</name>, 1923: Marine Littoral Plant and Animal Communities in New Zealand. <hi rend="i" TEIform="hi">Trans. Proc. N.Z. Instit.</hi> 54: 496-545.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-208922" reg="T. J. Parker" TEIform="name">Parker, T. J.</name>, 1886: Presidential Address to the Otago Institute. <hi rend="i" TEIform="hi">Trans. Proc. N.Z. Instit.</hi> 19: 613-615.</bibl>
<bibl default="NO" TEIform="bibl">Phillips, W. J., 1927: <hi rend="i" TEIform="hi">Bibliography of New Zealand Fishes.</hi> New Zealand Marine Department Bulletin No. 1, Wellington.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-208989" reg="J. S. Polack" TEIform="name">Polack, J. S.</name>, 1838: <title TEIform="title"><name key="name-134423" type="title" TEIform="name"><hi rend="i" TEIform="hi">New Zealand: Being a Narrative of Travels and Adventures During a Residence in that Country between the Years 1831 and 1837.</hi></name></title> Richard Bently, London.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-209010" reg="A. W. B. Powell" TEIform="name">Powell, A. W. B.</name>, 1937: New Species of Marine Molluscs from New Zealand. <hi rend="i" TEIform="hi">‘Discovery’ Rep.</hi> 15: 153-222.</bibl>
<bibl default="NO" TEIform="bibl">Richardson, J., 1843: Report on the Present State of the Ichthyology of New Zealand. <hi rend="i" TEIform="hi">Rep. Br. Ass. Advmt. Sci.</hi> 1843: 12-30.</bibl>
<bibl default="NO" TEIform="bibl">Sherrin, R. A. A., 1886: <hi rend="i" TEIform="hi">Handbook of the Fishes of New Zealand.</hi> Wilson and Horton, Auckland.</bibl>
<bibl default="NO" TEIform="bibl">Slack, E. B., <hi rend="i" TEIform="hi">et al.</hi>, 1969: <hi rend="i" TEIform="hi">Fisheries and New Zealand.</hi> Victoria University, Wellington.</bibl>
<pb id="n25" n="212" TEIform="pb"/>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-209371" reg="H. Suter" TEIform="name">Suter, H.</name>, 1913: <hi rend="i" TEIform="hi">Manual of New Zealand Mollusca.</hi> Government Printer, Wellington.</bibl>
<bibl default="NO" TEIform="bibl"><name type="person" key="name-209458" reg="G. M. Thomson" TEIform="name">Thomson, G. M.</name>, 1921: <hi rend="i" TEIform="hi">History of the Portobello Marine Fish Hatchery and Biological Station.</hi> Government Board of Science and Art, Wellington.</bibl>
<bibl default="NO" TEIform="bibl">Waite, E. R., <hi rend="i" TEIform="hi">et al.</hi>, 1909: Scientific Results of the New Zealand Government Trawling Expendition. 1907. <hi rend="i" TEIform="hi">Rec. Canterbury Mus.</hi> 1: 45-285.</bibl>
<bibl default="NO" TEIform="bibl">Watson, R. B., 1883: New Zealand Mollusca of the ‘Challenger’ Expedition. <hi rend="i" TEIform="hi">N.Z. Jl. Sci.</hi> 1: 319-321, 353-359, 441-443.</bibl>
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<bibl default="NO" TEIform="bibl"><name type="person" key="name-170474" reg="J. C. Yaldwyn" TEIform="name">Yaldwyn, J. C.</name>, 1957: A Review of Deep Water Biological Investigations in the New Zealand Area. <hi rend="i" TEIform="hi">N.Z. Sci. Rev.</hi> 15 (5-6): 41-45.</bibl>
</listBibl>
</div2>
</div1>
<pb id="n26" n="213" TEIform="pb"/>
<div1 id="t1-body-d2" type="article" decls="text-2-bibl" org="uniform" sample="complete" part="N" TEIform="div1">
<head TEIform="head"><title level="a" TEIform="title"><hi rend="b" TEIform="hi"><hi rend="c" TEIform="hi">Predators of Sea Anemones</hi></hi></title></head>
<byline TEIform="byline">by <name type="person" key="name-170576" TEIform="name">J. R. Ottaway</name><lb TEIform="lb"/>
Department of Zoology, University of Canterbury, Christchurch</byline>
<div2 id="t1-body-d2-d1" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Abstract</hi></head>
<p TEIform="p"><hi rend="b" TEIform="hi">A review is presented of literature concerning invertebrate and some vertebrate predators of sea anemones. Several previously unpublished records for New Zealand are given.</hi></p>
</div2>
<div2 id="t1-body-d2-d2" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Introduction</hi></head>
<p TEIform="p">Invertebrates, particularly the nudibranch <hi rend="i" TEIform="hi">Aeolidia papillosa</hi> (Boutan, 1898; Fleure and Walton, 1907), pycnogonids (Stephenson, 1928) and starfish (Verwey, 1930), have long been regarded as important or potentially important predators of sea anemones, and actual records are not uncommon for northern hemisphere species (Tables 1, 2).</p>
<p TEIform="p">There appears to be only one previously published record of an invertebrate preying on an anemone in the New Zealand and Australian region, and a few records for vertebrate predators (Table 3). This review therefore aims to encourage more records for these two countries.</p>
</div2>
<div2 id="t1-body-d2-d3" type="subsection" org="uniform" sample="complete" part="N" TEIform="div2">
<head TEIform="head"><hi rend="c" TEIform="hi">Discussion</hi></head>
<p TEIform="p">As predators of sea anemones, Stephenson (1928) cited ‘various fishes, crabs and other crustacea, nudibranchs, starfish, the larger worms [and] pycnogonids’. The relative importance of these predators is still difficult to assess, because of a lack of detailed field studies.</p>
<p TEIform="p">Aeolids have been the most frequently <hi rend="i" TEIform="hi">observed</hi> predators (Table 1), and Stephenson (1928) thought that these molluscs ‘are no doubt responsible for a good deal of damage’ to actinians in the wild. Fleure and Walton (1907) considered <hi rend="i" TEIform="hi">Aeolidia papillosa</hi> to be ‘the most formidable enemy sea-anemones possess’, and Harris (1971) suggested that seasonal fluctuations in the numbers of <hi rend="i" TEIform="hi">A. papillosa</hi> could influence the distribution patterns of its prey, <hi rend="i" TEIform="hi">Metridium senile.</hi> Certainly, this particular nudibranch exhibits a distinct preference for actinian species such as <hi rend="i" TEIform="hi">Actinia equina</hi> and <hi rend="i" TEIform="hi">Anthopleura elegantissima</hi> (Miller, 1961; Waters, 1973; Edmunds, Potts, Swinfen and Waters, 1975), to which it is attracted from some distance (Stehouwer, 1952; Braams and Geelen, 1953), but many more <hi rend="i" TEIform="hi">in situ</hi> studies, such as Wobber's (1970) observations of <hi rend="i" TEIform="hi">Dendronotus iris</hi> feeding on <hi rend="i" TEIform="hi">Cerianthus</hi> sp., are still needed.</p>
<p TEIform="p">The opisthobranch genus <hi rend="i" TEIform="hi">Aeolidia</hi> is poorly represented in the southern hemisphere, and apparently not at all in Australia or New Zealand (Robson, 1966); however, one might still expect to find a similar invertebrate predator filling the equivalent ecological niche. To date such predators have been largely undetected in this region, and little is known about the food sources of carnivorous opistho-branchs
<pb id="n27" n="214" TEIform="pb"/>
<table rows="45" cols="3" TEIform="table">
<head TEIform="head"><hi rend="c" TEIform="hi">Table 1. Records of Predation on Anthozoans by Aeolids in the Northern Hemisphere</hi></head>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell">PREY</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">PREDATOR</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">AUTHORITIES</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Actinia equina</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Boutan(1898); Stephenson (1928); Braams and Geelen (1953); den Hartog (1961); Miller (1961); Swennen (1961); Bruce, Colman and Jones (1963); Thompson (1964); Edmunds <hi rend="i" TEIform="hi">et al.</hi> (1974)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella sanguinea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Spurilla neapolitana</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Salvini-Plawen (1972)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Actinia sp.</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Fleure and Walton (1907); Eliot (1910)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Actinothoe anguicoma</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella glauca</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella sanguinea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aiptasia mutabilis</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Spurilla neapolitana</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Salvini-Plawen (1972)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anemonia sulcata</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Miller (1961); Edmunds <hi rend="i" TEIform="hi">et al.</hi> (1974)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Spurilla neapolitana</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Salvini-Plawen (1972)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthea sp</hi>.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Fleure and Wilton (1907); Eliot (1910)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthopleura artemisia</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Waters (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthopleura elegantissima</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Francis (1973); Waters (1973); Edmunds <hi rend="i" TEIform="hi">et al.</hi> (1974)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Hermissenda crassicornis</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Francis (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthopleura nigrescens</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Herviella</hi> sp. nov.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Rosin (1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthopleura xanthogrammica</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Waters (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Bunodeopsis</hi> sp.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Spurilla neapolitana</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Salvini-Plawen (1972)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Calliactis effoeta</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Boutan (1898)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Cerianthus</hi> sp.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Dendronotus iris</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Wobber (1970)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Cereus pedunculatus</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Boutan (1898); Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella alderi</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella glauca</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella sanguinea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965, 1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Cerberilla bernadetti</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965)</cell>
</row>
<pb id="n28" n="215" TEIform="pb"/>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Diadumene cincta</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Swennen (1961); Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella alderi</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella glauca</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella sanguinea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965, 1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Cerberilla bernadetti</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Diadumene luciae</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Waters (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Epiactis prolifera</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Waters (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Metridium marginatum</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Russell (1942)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Metridium senile</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Stehouwer (1952); Braams and Geelen (1953); Swennen (1961); Wolter (1967); Harris (1971)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Sagartia elegans</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella glauca</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Miller (1961); Bruce, Colman and Jones (1963)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Sagartia troglodytes</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Miller (1961); Swennen (1961); Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella glauca</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965, 1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella sanguinea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965, 1969)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Cerberilla bernadetti</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Tardy (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Sagartia</hi> sp.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidiella takanosimensis</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Salvini-Plawen (1972)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Berghia coerulescens</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Salvini-Plawen (1972)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Stomphia coccinea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Robson (1961)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia glauca</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Miller (1961)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Tealia coriacea</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Waters (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Tealia crassicornis</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">McMillan (1942); Waters (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Tealia felina</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Aeolidia papillosa</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Miller (1961); Swennen (1961)</cell>
</row>
</table>
<table rows="28" cols="3" TEIform="table">
<head TEIform="head"><hi rend="c" TEIform="hi">Table 2. Other Records of Predation on Anthozoans in the Northern Hemisphere</hi><note id="fn1-215" n="*" place="unspecified" anchored="yes" TEIform="note"><p TEIform="p">Further references to associations of pycnogonids and anthozoans are listed in Helfer and Schlottke (1935), Hedgpeth (1949), and Wyer and King (1974).</p></note></head>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="c" TEIform="hi">Prey</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="c" TEIform="hi">Predator</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="c" TEIform="hi">Authorities</hi></cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Actinia equina</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Nymphon gracile</hi> (Pycnogonida: Nymphonidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Wyer and King (1974)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Pycnogonum littorale (Pycnogonida: Pycnogonidae)</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Bruce, Colman and Jones (1963); Myer and Buckmann (1963); Wyer and King (1974)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anemonia sulcata</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Pycnogonum littorale</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Wyer and King (1974)</cell>
</row>
<pb id="n29" n="216" TEIform="pb"/>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthopleura elegantissima</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Calliostoma annulatum</hi> (Gastropoda: Trochidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Francis (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Dermasterias imbricata</hi> (Asteroidea: Poraniidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Mauzey, Birkeland and Dayton (1968)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Epitonium</hi> sp. (Gastropoda: Epitoniidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Francis (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Patiria miniata</hi> (Asteroidea: Asterinidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Francis (1973)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Pycnogonum stearnsi</hi> (Pycnogonida: Pycnogonidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Fry (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Anthopleura xanthogrammica</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Dermasterias imbricata</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Mauzey <hi rend="i" TEIform="hi">et al.</hi> (1968)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Pycnogonum stearnsi</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Fry (1965)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Calliactis parasitica</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Crossaster papposus</hi> (Asteroidea: Solasteridae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Milligan (1916)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Pycnogonum littorale</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Wyer and King (1974)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Diadumene</hi> sp.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Crossaster papposus</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Hancock (1958)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Edwardsia</hi> sp.</cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Platichthys flesus</hi> (Pisces: Pleuronectidae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">McIntosh (1874)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Epiactis prolifera</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Dermasterias imbricata</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Mauzey <hi rend="i" TEIform="hi">et al.</hi> (1968)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Metridium senile</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Dermasterias imbricata</hi></cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Mauzey <hi rend="i" TEIform="hi">et al.</hi> (1968)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Gephyreaster swifti</hi> (Asteroidea: Radiasteridae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Mauzey <hi rend="i" TEIform="hi">et al.</hi> (1968)</cell>
</row>
<row role="data" TEIform="row">
<cell role="data" rows="1" cols="1" TEIform="cell"/>
<cell role="data" rows="1" cols="1" TEIform="cell"><hi rend="i" TEIform="hi">Hippasteria spinosa</hi> (Asteroidea: Goniasteridae)</cell>
<cell role="data" rows="1" cols="1" TEIform="cell">Mauzey <hi rend="i" TEIform="hi">et al.</hi> (1968)</cell>
</row>
<row role="data" TEIform="row"