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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
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).
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).
After Tasman's departure, more than 125 years were to pass before another European expedition came to New Zealand. The arrival of Captain
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Hawkesworth was published in 1773, but Cook's journal itself was not printed until 1893, and Banks's journal remained unpublished until it appeared in a considerably edited and modified version in 1896.
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
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 Systema Ichthyologiae, 1801, but the complete systematic account of the scientific results of the voyage, written by
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
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
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, et al., 1953).
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).
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
Memoire Geographique sur la Nouvelle Zelande published in 1826 (McNab, 1907).
Perhaps the one ship that contributed the most to New Zealand marine biology in this period was a French corvette, originally named Coquille, 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.
In 1827 the Coquille, now renamed Astrolabe, 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
The Astrolabe made her third and final stop in New Zealand in 1840 on a voyage to the Antarctic, this time accompanied by another corvette, Zelee. 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 et al., 1953).
During the period from 1835 to 1850, four of the most famous naturalists of the nineteenth century visited New Zealand.
Darwin visited the Bay of Islands for nine days in December 1835 while serving as naturalist on H.M.S. Beagle 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).
Hooker accompanied James Clark Ross on the vessels Erebus and Terror 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 et al., 1953). The Erebus and Terror 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.
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.
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.
Huxley spent a week in the Bay of Islands in May 1850 while H.M.S. Rattlesnake, on which he was serving as assistant surgeon,
During the years 1847 to 1849, H.M.S. Acheron 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 Acheron, 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.
The last noteworthy expedition of this era was that of the Austrian ship, Novara, which visited Auckland and the Novara's 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.
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.
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.
The first European resident to provide information on marine biology was
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.
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 Travels in New Zealand in 1843.
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.
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
At this time the need for a national scientific body was realised, and in 1867
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).
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 Transactions and Proceedings 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).
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
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.
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,
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,
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.
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
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, The Fishes of New Zealand in which he described the 141 species of fish known at that time to exist in New Zealand waters. This was followed by the Catalogue of Echinoderms in New Zealand, 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 Catalogue of the Marine Mollusca of New Zealand 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 Index Faunae Novae Zelandiae, published in 1904.
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
No description of the history of marine biology in any part of the globe would be complete without some mention of the voyage of H.M.S. Challenger. Many people today consider the Challenger 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 Challenger 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 Challenger 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 et al., 1895).
The Challenger 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 Challenger 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).
At the same time as the Challenger 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.
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 Erebus and Terror 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
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. Challenger, 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 Transactions and Proceedings of the New Zealand Institute for these years contain numerous reports of their findings in marine biology.
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.
The need for a marine station was first noted by
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.
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).
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.
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 N.Z. G. S. Stella 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
S.S. Plucky. 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).
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
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.
The Philosophical Institute of Canterbury organised an expedition to the Auckland and Campbell Islands in November 1907 on the government steamer Hinemoa. 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
In 1908 a private party on the steamer Rakiura 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.
The early part of the twentieth century saw steady progress being made by New Zealanders in almost all areas of marine biological
Astrolobe, conchology had always been at the forefront of marine science in New Zealand, and this trend continued after 1900. Manual of New Zealand Mollusca. 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.
Similar, though less spectacular, advances were being made in the studies of other groups of marine animals at this time.
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 Marine Littoral Plant and Animal Communities in New Zealand, published in 1923.
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 Challenger 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.
The first of these visitors was the British Terra Nova 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’ (British Antarctic Expedition, 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.
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 Hinemoa 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 G.S. Amokura 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 Videnskadelige Meddelelsen fra Danske Naturhistorrisk Forering i Kobenhavn throughout the 1920's.
Another Danish expedition, the Carlsberg Foundation Oceanographic Expedition around the world 1928-30 on the Dana, arrived in Auckland in December 1928. From there the Dana 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 Dana Reports in Copenhagen in the 1930's.
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.
British scientists returned to New Zealand in August 1932 on the R.R.S. Discovery II. They took new samples from some of the bottom stations examined by the Terra Nova and investigated some additional areas suggested by Terra Nova, the Discovery 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.
Throughout its history, New Zealand marine biology has been strongly affected by the fisheries. The desire to capture food from the
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,
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 Doto 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 Doto and then the Rita 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 et al., 1909).
As a follow-up to the 1900-01 expedition, the government chartered the Nora Niven 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
et al., 1909). As late as 1947, Waite's report on the Nora Niven expedition was still considered to ‘represent the most complete and comprehensive monograph on New Zealand ichthyology yet published’ (Hefford, 1947).
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).
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.
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 Discovery II on a voyage to Antarctica and gained valuable training in modern methods of biological investigation.
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,
In the absence of a research staff suitably equipped with laboratory accommodation and with facilities for pursuing
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 Ikatere 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.
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.
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.
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.
In 1946 the D.S.I.R. sponsored an expedition to the southern fiords on the New Golden Hind. During the expedition 84 bottom samples were taken with a Peterson grab from depths of two to 73 fathoms.
Much material of interest was found in these samples, including 342 species of molluscs, 17 of which were new (Fleming, 1950).
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 H.M.S. Lachlan 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 Lachlan and arranged for the examination of all the specimens taken by the vessel.
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).
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.
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
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.
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 New Zealand Journal of Marine and Freshwater Research in 1967 has added yet another channel of communication for marine biologists.
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.
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.
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
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
A review is presented of literature concerning invertebrate and some vertebrate predators of sea anemones. Several previously unpublished records for New Zealand are given.
Invertebrates, particularly the nudibranch Aeolidia papillosa (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).
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.
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.
Aeolids have been the most frequently observed 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 Aeolidia papillosa to be ‘the most formidable enemy sea-anemones possess’, and Harris (1971) suggested that seasonal fluctuations in the numbers of A. papillosa could influence the distribution patterns of its prey, Metridium senile. Certainly, this particular nudibranch exhibits a distinct preference for actinian species such as Actinia equina and Anthopleura elegantissima (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 in situ studies, such as Wobber's (1970) observations of Dendronotus iris feeding on Cerianthus sp., are still needed.
The opisthobranch genus Aeolidia 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
Few other molluscs have been reported as predators of anemones (Tables 2, 3). Francis (1973) recorded Calliostoma annulatum and Epitonium sp. as predators of Anthopleura elegantissima, but gave no indication of their effect on populations. Fleure and Walton (1907) ‘once noticed Trochus ziziphinus nibbling at the base of an anemone’, and on two occasions I observed Haustrum haustorium to partially eat juvenile Actinia tenebrosa. The two juveniles died as a consequence of the attacks, but it seems unlikely that predation by Haustrum would have any marked effect on Actinia populations.
The polychaete worm Hermodice carunculata is known to be an active predator of Stoichactis helianthus. After a single Hermodice starts feeding on Stoichactis, other worms are attracted to feed on the same anemone. Although it is not known whether the anemones die following such attacks, Hermodice could still affect the local distribution of Stoichactis (Lizama and Blanquet, 1975). A similar situation might exist for pycnogonids. Even though Fry (1965) considered actinians to be the major food source of Pycnogonum stearnsi, it is not known whether predation by sea-spiders would actually kill adult anemones. Stephenson (1928), however, observed that ‘serious results may ensue from such attacks on young or small anemones’, so there is still the possibility that pycnogonids could influence the distribution of some actinian species.
Some starfish are considered to be omnivorous (Feder and Christensen, 1966) and have been observed to eat anemones in an aquarium (Milligan, 1916) and in the sea (Table 2). Verwey (1930) believed that Acanthaster echinites was ‘probably one of the worst enemies of anemones’; however, the first unequivocal indication that actinians can be a significant part of the diet of some asteroids came from the extensive in situ observations of Mauzey, Birkeland and Dayton (1968). They found that, in some habitats, Dermasterias imbricata fed mainly on Epiactis prolifera, and occasionally on Anthopleura xanthogrammica, A. elegantissima, Tealia coriacea and Metridium senile. Other starfish were also seen to eat anemones (Table 2).
Most known vertebrate predators are fish. Over a century ago Peachia hastata had been discovered in cod stomachs, and ‘swarms’
Edwardsia species in flounder stomachs (McIntosh, 1874). Even within symbiotic relationships, damselfish sometimes eat the tentacles of their host anemone, Stoichactis (Verwey, 1930). Several New Zealand fish species ingest anemones (Table 3), although from the two detailed records it seems likely that the anemones may have been ingested incidentally along with other prey. Webb (1973) found that an Anthopleura species constituted up to 4.8% of the gut contents of the sand flounder Rhombosolea plebeia, but the cockle Chione stutchburyi is a significant part of the diet of this flounder, and where the study was done Anthopleura aureoradiata is commonly found attached to the cockle (Parry, 1951). Similarly, 40 Calliactis conchicola and several Paracalliactis rosea were seen in the gut contents of 12 gummy sharks, Emmissola antarctica (= Mustelus antarcticus), caught in 100m of water offshore from Kaikoura (Ottaway, personal observation). At Kaikoura, both anemones are epizoic on the carapaces of the spider crab, Leptomithrax longipipes, and C. conchicola is also found on living whelks, Austrofusus glans, and whelk shells occupied by hermit crabs (Hand, 1975). Every anemone found eaten by the gummy sharks was attached to Leptomithrax, but there were Leptomithrax present in the stomachs that did not bear anemones. Since there were no whelks or hermit crabs ingested, with or without anemones, it seems that the sharks were preying on Leptomithrax rather than the anemones.
Other vertebrate predators seem to be few. At one time the French collected anemones and sold them in Bordeaux markets (Rondeletius, quoted in Johnstone, 1846, p. 227). Dicquemare (1773) actually carried out cautious experiments on the toxicity of cooked actinians, feeding them to his cat and himself. He apparently satisfied himself that some species, especially Tealia crassicornis, were palatable, because later he wrote (Dicquemare, 1775): ‘De toutes les efpeces d'anémones de mer, celle-ci m'a paru devoir mériter la préférence pour la table. Lorfqu'on les a fait bouillir un peu fermes, et qu'on les fert pour manger à quelle fauffe on juge àpropos, …’ Even so, Johnston (1846) had reservations about Dicquemare's opinion: ‘The mouth waters at the liquorish description, … but I have not been tempted to test its truth.’ More recently, Martin (1960) reported that cooked Rhodactis howesii was commonly eaten by Samoans, although fatal cases of poisoning were known to occur amongst those natives who had eaten the anemone raw.
I am indebted to Dr W. C. Clark, Professor
This work was supported by a New Zealand Commonwealth Postgraduate Award.
Riccia ciliata, R. crozalsii, R. glauca, R. sorocarpa and R. bullosa are recorded for New Zealand. A key is given to the Riccia species known to be growing in New Zealand at present.
Since the publication of an earlier paper (Campbell, 1975), five additional species of Riccia have been found in New Zealand. Of the four which belong to the subgenus Riccia, two possess cilia and two lack this feature. The fifth species is placed in the subgenus Ricciella, although the structure of its thallus is intermediate between the two subgenera. Notes on the five species are given below.
Description of the Plant
The plants are found on soil as greyish-green or glassy-green rosettes up to 3cm in diameter (Fig. 1). Branches are up to 3mm long and 0.6-0.8mm wide. The upper surface of the thallus is flat or convex except for a narrow median groove near the apex. Under a magnification of X 10 or higher it appears marked out into squares. Acicular spines, 0.6-0.9mm long and with a basal width of 40-60μm. project from the margins. They have thick walls which, when swollen with water, tend to be internally tuberculate. On the under side of the thallus there are hyaline median scales, smooth rhizoids of diameter 18-30μm and tuberculate rhizoids of diameter 10-18μm.
Plants are monoicous. The spores have plate-like reticulate markings along with numerous papillae. They have a diameter of 70-90μm, occasionally up to 110μm (Fig. 2a and b).
Anatomy of the Thallus
In transverse section the thallus is approximately the same depth of 0.3-0.4mm throughout. The epidermis consists of hyaline, thin-walled cells which at first are of spherical or conical shape but later shrivel and flatten. The photosynthetic tissue is composed of columns of cells separated by vertical air canals, and beneath is a compact tissue of hyaline cells.
Distribution
R. ciliata is widely distributed in Europe. Probably it has been introduced to New Zealand.
It was collected from seepage areas on the shore of Lake Pukaki where it was growing with other species of Riccia,
Description of the Plant
The plants are found on soil as firm, green rosettes up to 1.5cm in diameter or as large irregular colonies (Fig. 3). Branches are up to 4mm long and up to 1 mm wide. The thallus at a magnification of X 10 or higher often appears spongy due to the spherical, hyaline, epidermal cells (Fig. 4). In older parts it has a flat or slightly convex surface, but near the apex it rises steeply on either side of a well-defined narrow median groove. Hyaline, sharp-pointed cilia, up to 0.5mm long, project from the margin or incurve over the thallus, but they often wither early and are invisible when plants are flooded with water. Their surface, particularly in the upper part, is minutely tuberculate. On the under side of the thallus are smooth rhizoids of diameter 8-21μm, tuberculate rhizoids of diameter 10-18μm and small, delicate hyaline scales. Branching is by forking at the apex and occasionally by adventitious branches.
Plants are monoicous. The dark brown spores have triradiate and reticulate markings and truncate, papillate projections. There are conspicuous germ-pores and a broad wing (Fig. 5).
Anatomy of the Thallus
The thallus in transverse section is 0.4mm deep. The dorsal epidermis consists of more or less spherical, hyaline cells. A few of the lowermost cell layers are also colourless. The rest of the thallus is a rather compact, photosynthetic tissue of subspherical cells with air spaces between them.
Distribution
R. crozalsii is known from the western part of Europe, the Canary Islands, and the Cape Province of South Africa (Arnell, 1963); from Israel (Proskauer, 1953); from South India (Udar, 1957) and from South Australia and New South Wales (Seppelt, 1974). In New Zealand it is very plentiful near paths on Rangitoto Island, Auckland, Riccia species on the summit of Mt. Eden, J. E. Braggins,
Notes
Some forms of R. crozalsii are not readily distinguishable from R. bifurca and the spore morphology of these two species corresponds closely. A detailed study has been made by Seppelt (1974).
Description of the Plant
The plants grow on soil as glaucous-green rosettes up to 3cm in diameter or as irregular colonies (Fig. 6). Branches are strap-shaped, up to 10mm long and 1-1.5mm wide. The upper surface of the thallus is flat in older parts but near the apex has a broad, shallow median groove. In young parts it has a glistening appearance and later shows a fine polygonal pattern (Fig. 7). On the under side of the thallus are delicate, hyaline scales which reach to the margin and both smooth and tuberculate rhizoids.
The plants are monoicous. The spores, of diameter 75-90μm, have a broad wing and show reticulate markings with papillate projections on the distal face (Fig. 8).
Anatomy of the Thallus
The thallus in transverse section is 0.3-0.6mm deep. The dorsal epidermis consists of either one or two layers of thin-walled, hyaline cells. Those of the outermost layer vary from pear-shaped to rounded and tend to collapse early. Those of the underlying layer persist but do not become thick-walled as in R. sorocarpa. Beneath the epidermis is a photosynthetic tissue, which consists of columns of cells enclosing narrow, vertical air canals, while the lowermost third of the thallus is a compact, hyaline tissue.
R. glauca is found in Europe, America, Asia and Africa. Watson (1959) states that it is perhaps the commonest British Riccia species. Probably it has been introduced into New Zealand.
It was collected from seepage areas on the shores of Lake Pukaki, growing amongst other Riccia species,
Description of the Plant
The plants occur on soil, usually as glaucous-green rosettes or partial rosettes which are 4-9mm in diameter and 2-4 times dichotomously branched. Occasionally there are irregular colonies. Branches are 1-2mm broad. They are narrowed towards the subacute apex and here show a deep and narrow median furrow which becomes less pronounced in older parts. At a magnification of X 10 or higher the upper surface in young parts appears glistening after the manner of R. crystallina. In older parts, after the outermost cells have been destroyed, the surface appears marked into squares. On the under side there are small, hyaline scales which reach to the margin and both smooth and tuberculate rhizoids. Plants are monoicous. The spores are about 80μm in diameter. On the distal face they show reticulate markings with high papillae in the corners (Fig. 9) and on the proximal face they show short, low ridges and a triradiate marking.
Anatomy of the Thallus
The thallus is 0.3-0.6mm. deep. The dorsal epidermis, which provides a useful diagnostic feature for this species, consists of two layers of hyaline cells. The conical or pear-shaped cells of the outer layer are soon destroyed, leaving intact the underlying layer of thick-walled cubical cells (Fig. 10a and b). The lower half of the thallus consists of thin-walled hyaline cells arranged compactly. The rest is a photosynthetic tissue in the form of columns of cells enclosing narrow air canals.
Distribution
R. sorocarpa is widely distributed. It is well known in Europe, North America, Japan, Siberia, North and South Africa (Arnell, 1963). It is reported also from South America (Hassel de Menendez, 1962), from the Sikim Himalayas (Udar, 1956) and from Australia (Seppelt, 1974). In New Zealand it has been found growing with other Riccia species in seepage areas on the shore of Lake Pukaki,
Notes
R. sorocarpa may be indigenous to New Zealand, for Mitten (1855) recorded R. acuminata Tayl, as collected by Colenso in
R. acuminata (Melb. 19974) was examined by O. Na-Thalang (1970) and by R. D. Seppelt who found that it contained both R. sorocarpa and R. lamellosa. Seppelt notes that, since Taylor's description (1846) is based on mixed material, the name R. acuminata becomes a nomen confusum and by Art. 70 of the International Code of Botanical Nomenclature must be rejected (Seppelt, 1974).
In the Colenso Herbarium of the National Museum of New Zealand there is a specimen which is almost certainly a duplicate of the R. acuminata Tayl. listed by Mitten (1855). This specimen was examined superficially and is considered to be R. sorocarpa. Despite a thorough search in the spring of 1974 no plants of Riccia were found in Hawke's Bay.
Description of the Plant
The plants are found on soil, usually as strap-shaped thalli up to 13mm long and up to 4mm wide, once or twice forked; occasionally as rosettes, partial rosettes or large irregular colonies. The colour is green or olive-green above and either green or reddish-purple below. The upper surface is generally flat or convex, with a deep, narrow median groove near the apex, and has thin edges which tend to be hyaline and crenate. Under a magnification of X 10 or higher it is seen to be marked out into polygonal areas whose boundaries represent partitions between sub-epidermal air cavities (Fig. 11). Air pores show clearly in scanning electron micrographs (Fig. 12). The under surface carries both smooth and tuberculate rhizoids, and near the apex a few delicate, hyaline scales which reach to the thallus edge. However, sometimes, as mentioned later, scales do not develop.
In regard to their sexual nature some plants are definitely monoicous, in others only antheridia or only archegonia are present, while many plants are sterile. The brown spores, 0.08-0.1mm in diameter, have a crenate wing and show reticulate and a faint triradiate marking. The areolae on the distal face have a thin border with higher and thicker ridges at the corners; those on the proximal face are ill-defined (Fig. 13a and b).
Anatomy of the Thallus
The thallus shows differentiation into epidermis, spongy tissue. and compact ventral tissue. In transverse section it is 0.6-0.8mm deep in the central region but gradually narrows laterally to a wing-like edge only one cell deep (Fig. 14). The cells of the upper epidermis tend to be hyaline around the air pores. Beneath is a spongy photosynthetic tissue, which in the central region is 0.15mm deep and is composed of vertical partitions separating narrow air canals (Fig. 15), and nearer the margin is 0.3-0.45mm deep and consists of unistratose partitions between what appear as two or three layers of wider air cavities (Fig. 16). The ventral tissue, 0.2-0.4mm deep, is green in young plants of the thallus but has only a few chloroplasts in older parts.
Distribution
Riccia bullosa is found in South Africa (Sim, 1926; Arnell, 1963). In the North Island of New Zealand it has been collected in the
Notes
As a result of a comparison between herbarium specimens of R. bullosa from South Africa and plants collected in New Zealand it was concluded that the species exhibits a considerable degree of polymorphism. Plants from South Africa tended to be firmer due to epidermal cells with somewhat thickened walls. In plants from the South Island of New Zealand the air pores were less clearly defined than in those from the North Island and the cells surrounding them tended to disintegrate in older parts of the thallus. Also scales failed to develop. Lack of scales was found also in plants collected in the North Island and grown in culture under conditions of high humidity and cool temperature.
As was noted by Arnell (1963), the structure of the thallus in R. bullosa is intermediate between that of the subgenus Riccia and the subgenus Ricciella. There are narrow air canals in the central region and much wider air cavities nearer the margin. This type of structure is found also in R. vesiculosa Carr. and Pears., as noted by Stephani (1898-1900). The range of R. vesiculosa is given as New South Wales, Queensland, Northern Territory (Na-Thalang, 1970).
Carrington and Pearson (1887) mention the close similarity of R. vesiculosa to R. bullosa but the latter is stated to be dioicous (Sim, 1926; Arnell, 1963) in contrast to the reported monoicous condition of R. vesiculosa (Stephani, 1898-1900; Na-Thalang, 1970). The sexual nature of the New Zealand plants was found to be very variable.
The writer is indebted to R. D. Seppelt for identifying specimens of R. crozalsii and R. sorocarpa and for discussion regarding the other species, to Prof. E. A. Schelpe for sending herbarium specimens of R. bullosa from South Africa, to
The first part of this series (McQueen, 1976) dealt, briefly, with the distribution and general characters of the ten species of Nothofagus in South America; and in more detail with the vegetation of the Magellanic region around latitude 52° to 55° S. A paper by Weinberger (1973), translated in this issue, deals bioclimatologically with the commoner species of Nothofagus, mostly around latitude 38° - 41°. The present contribution aims to complement and introduce his work, particularly from a New Zealand point of view. Comments are drawn from field observations in Chile early in 1972, and from a selection of the literature. A notable and well illustrated background to Chilean vegetation is di Castri (1968).
A general diagram of the relation of Nothofagus to altitude and precipitation around latitude 40° S. (Table 1) is repeated here for convenience from Part I (McQueen, 1976). Species of wide distribution in mid-latitude Chile are N. obliqua, N. antarctica, N. alpina (syn.
Due to a typographical error N. alpina was wrongly shown as being in the N. ‘fusca’ pollen group in Table 1 (McQueen 1976). It is, of course, in the N. ‘menziesii’ group.
To the New Zealander, Chilean forests around latitude 40° S. present some familiarities and many more unfamiliarities; this is a character already outlined by Godley (1960). There are both Nothofagus-dominated forests and dicotylous Nothofagus-absent forests in both countries. Sharing Podocarpus, Nothofagus, Weinmannia, Laurelia, Pseudopanax, Aristotelia and thirty-nine families with New Zealand the lowland Chilean forests are different, in the relation of species composition to structure, from New Zealand forests. One most notable difference is the understorey of the bamboo, Chusquea, in forests up to about 1,000 m altitude. Its density, in the dozen or so species found, is an important factor in prevention of regeneration when forest management is practised.
It appears that Nothofagus in South America is at a competitive advantage over the majority of Chilean conifers. Reasons may be found in its biological characters, such as deciduousness, and the ability of some species to regenerate vegetatively, and apparently easier seed dispersal (McQueen, 1976). There is a marked contrast, at least on younger soils, with New Zealand, in the relation between the two groups of trees. Chilean Podocarpus plays a sub-canopy role in Nothofagus forest and dicotylous forests. Saxegothea conspicua, in the same family, will reach the upper canopy level with Nothofagus and occasionally will form pure stands (Schmithusen, 1960). Thus S. conspicua is the only tree equivalent in its structural position as a canopy or emergent tree to New Zealand species of Podocarpus and Dacrydium. The only representative of Dacrydium in Chile is a prostrate swamp plant slightly bigger than Dacrydium laxifolium. It is, however, ecologically equivalent to Dacrydium bidwillii.
Two members of the Cupressaceae in Chile are Pilgerodendron uviferum and Fitzroya cupressoides, both found growing in poorly drained soils and even in peat swamps. The first is restricted to such sites and extends southwards to the western part of the Magellanic region (McQueen, 1976). The second species, F. cupressoides, does not grow as far south, but as well as growing in bogs it will grow on very infertile soils at altitudes up to 1,000 m on the Coastal Range. These two species behave similarly to Dacrydium colensoi and Dacrydium intermedium in New Zealand. Also in the Cupressaceae in Chile is Austrocedrus (formerly Libocedrus) with one species, A. chilensis, whose ecology is different from its New Zealand relatives (L. bidwillii, L. plumosa), both of high precipitation forest areas. The Chilean species is a plant growing to the north of Nothofagus forest on the Andes as rainfall decreases northwards. It also occurs in Argentina in the rain shadow produced by the Andes mountains, where it is accompanied by N. antarctica, the most drought tolerant of South American Nothofagus (Fig. 1). A. chilensis also occupies drier intermontane basins in Chile, west of the Andes.
A similar contrast is found in the Araucariaceae: New Zealand's Agathis australis is northern, almost sub-tropical; Chile's
Hybridisation may be as prevalent among Nothofagus in South America as in New Zealand. Forms seen by the author included a N. obliqua with N. glauca characters, at the north of the Nahuelbuta National Park at 37° 47′ S, and the apparent variety N. obliqua var. macrocarpa of the Cerro Roble at 33° S. Even the variety in leaf size of N. obliqua s.s. collected further south suggests a hybridism in two representatives of the N. ‘menziesii’ pollen group in South America. By contrast, in the N. ‘fusca’ group representatives, among the deciduous species the common, and often adjacent, N. antarctica and N. pumilio retain their own characters. Amongst the evergreens of
N. dombeyi and N. betuloides, although very similar, and overlapping slightly in range at about 40° - 41° S, are always clearly distinguishable.
Returning to characters of the South American representatives of the N. ‘menziesii’ group: N. obliqua and N. glauca, these deciduous species have a different ecology from the evergreen N. menziesii of New Zealand. N. obliqua and N. glauca are lowland, and northern in distribution, tolerating a degree of summer drought.
The implications to paleoecology of the N. ‘menziesii’ pollen group are important. This pollen has existed in New Zealand since the Cretaceous (Couper, 1960) and has been interpreted in pre-Pleistocene times as an indicator of cooler climates (McQueen et al., 1968). Such opinions may need revising if this group were, in the Cretaceous and Tertiary, represented in New Zealand by more thermophilic species of the group, such as exist today in South America and Australia.
The following notes are a commentary on the translation of Weinberger's paper of 1973 on page 245 of this issue of Tuatara, and follow his order of description.
Nothofagus obliqua
Nothofagus obliqua (Fig. 3) is deciduous and is capable of vegetative reproduction after damage by fire. It was formerly widespread in the central valley of Chile from 38° S near sea level to about 41° S (Clarke, 1964). N. obliqua does extend further north at increasing altitude and is the northernmost Nothofagus found in Chile, recorded in its form N. obliqua var. macrocarpa as far north as the Cerro Roble (33° S), part of the Coastal Range above the truly Mediterranean climate of Central Chile. At this locality the present author saw only its lower limits, very deformed trees in gullies forming tongues down into the Mediterranean scrub. This scrub contained mostly species quite unknown in New Zealand, but did include, however, Sophora macrocarpa and Muehlenbeckia hastata.
In its higher altitude northern distribution at about 700 m to 800 m Nothofagus obliqua exists as a distinct ‘form’ recognised by Chilean
N. obliqua, up to 1,350 m altitude on the same range, were growing on flat ground, frequently frosty, as described by Weinberger. The inclusion of this area within the National Park and protection from grazing had resulted in very good regeneration by seed.
It is in its main lowland distribution from 38° - 41° S that N. obliqua attains its best growth, up to 40m height. As shown in Weinberger (Fig. 5) N. obliqua ascends only to about 500m altitude on the drier eastern flanks of the coastal range at latitude 40° S. It is found to a similar altitude in the Andes where it frequently grades into Valdivian rainforest with N. alpina as an emergent.
In the Central Valley most of the forests of N. obliqua have been converted to farm land but the vegetative regrowth of N. obliqua has allowed scattered trees to remain, giving a very European appearance to the landscape.
The occurrence of summer frosts on the plains occupied by Nothofagus obliqua is ample evidence of cold air drainage off the
N. obliqua as a deciduous tree is well suited to such sites and to the higher altitude flat sites described by Weinberger.
Edaphically (Wright, 1965), N. obliqua is associated with the trumao soils, formed from volcanic ash with allophane as a dominant clay. These soils are characterised by friability to depth and high water retention throughout but nevertheless only a limited depth is penetrated by tree roots.
In the Central Valley the contrasting poorly drained, reworked volcanic ash nadi soils are the site for trees of lower fertility demand such as Nothofagus nitida, and formerly stands of very large Fitzroya cupressoides.
Nothofagus antarctica
Nothofagus antarctica rarely grows to more than 10m high and is multi-branched, deciduous as it N. obliqua. As described by Weinberger, it behaves in a similarly plastic way in middle latitude
nadi soils (Wright, 1965). It descends in cold air drainage sites to 900m or less, and as in the far south forms a subalpine scrub above the N. pumilio treeline on both flanks of the Andes, frequently in association with Araucaria araucana on the Andes and on the Coastal Range (Fig. 4). Weinberger rightly considers N. antarctica as a plant with a wide tolerance of thermal and atmospheric saturation ranges. He further draws a very interesting comparison involving the occurrences of N. antarctica in the Andes, where it occupies valley flats above about 900m. Here the valley floor is occupied by N. antarctica, and other Nothofagus species are located in the zone above the occurrence of heavy frosts. Weinberger points out that similar valley bottom sites in Magellanic Chile would be occupied by N. pumilio. The general temperature regime of the Magellanic area is considerably colder than that of the Andes, particularly during the growing season. Weinberger suggests that the virtually constant westerly winds of the Magellanic area cause far greater mixing of air masses and prevent cold air
N. pumilio on many of the flat areas around Punta Arenas. However, the present author's observations around Puerto Natales (latitude 52° S), where there are extensive areas of N. antarctica in valley bottom flats with N. pumilio on the slopes, suggests that further away from the more oceanic conditions around the Straits of Magellan, summer frost may play an equally important role in the distribution of extensive stands of N. antarctica in valley bottoms in the Magellanic area.
Nothofagus alpina
Nothofagus alpina (Fig. 5), which occurs above N. obliqua on the Coastal Range and the Andes, is likewise deciduous, intermingling with N. obliqua lower down and with the evergreen N. dombeyi above. In common with N. obliqua, N. alpina has vigorous vegetative reproductive habits and one stand seen in the Andes originated from trees engulfed in 60cm of basaltic ash in about 1870. It then resprouted, was felled about 1945, and resprouted again. Excavation showed that the regeneration was from the root collar and from root suckers.
Nothofagus pumilio
Nothofagus pumilio, as in Magellanic Chile, forms the treeline on the Andes, the small-crowned deciduous trees looking, in leaf,
N. solandri var, cliffortioides (Fig. 6). The forest floor at altitude is generally herbaceous and open but at lower altitudes in the N. pumilio zone the bamboos, Chusquea spp., are, to New Zealand eyes, odd companions to Nothofagus (Fig. 7).
It is in the upper parts of the N. pumilio zone that Araucaria araucana occurs in large but rather isolated areas between 38° and 40° S on the Andes and in the Nehuelbuta region of the Coastal Range at about latitude 37° S. This erect-growing conifer is either emergent above the N. pumilio canopy or above a matrix of low N. antarctica scrub above the N. pumilio treeline (Fig. 2). Apparently the robust yet flexible branches of Araucaria shed the snow that is the cause of the treeline krummholz of N. pumilio (Eskuche, 1973), thus giving a double treeline.
Nothofagus dombeyi
N. dombeyi (Fig. 8) is a tree climatically of a very wide tolerance, concentrated in Chile in climates of oceanic tendency, but widespread there on to sites of edaphic stress, such as the nadi soils, moraines and lahars. It will thrive as a coloniser even on such ‘new’ sites in areas exposed to the dry easterly wind, the puelche, whose temperature extremes outdo most Canterbury nor'-westers (see Fig. 8 of Weinberger).
It is in Argentina that N. dombeyi reaches its fullest and most continuous extent. On the rain shadow eastern flank of the Andes this species descends from the sub-alpine N. pumilio forests down to the semi-arid N. antarctica and Austrocedrus chilensis forests of the western Patagonian plains.
In its tolerance of the rain shadow climate of the east of the Andes and in many of its attributes in the higher rainfall areas of Chile N. dombeyi appears remarkably similar in its ecology to N. solandri var. cliffortioides, of New Zealand, but N. dombeyi does not ascend to the treeline. At its easternmost limit of precipitation in Argentina N. dombeyi is found only on relatively deep and friable volcanic ash soil. Rocks from which the ash has been eroded and recent shingle fans are occupied by Austrocedrus chilensis, which as well grows erect, beyond any continuous cover of N. antarctica.
Nothofagus nitida
Nothofagus nitida (Fig. 9), the other evergreen species treated in detail by Weinberger, is found only on the Coastal Range and to the south of Puerto Montt in the areas where the chain of the Andes is near the sea. N. nitida is a species of quite narrow ecological tolerance to temperature and moisture fluctuations. It is most similar in its growth and in its favoured habitats to Nothofagus menziesii in New Zealand, but does not reach the alpine treeline.
Although Weinberger had only one meteorological station for N. betuloides, it may be considered that N. betuloides, being predominant
N. nitida. They both occupy very similar climatic conditions.
There remain three species not studied in detail by Weinberger; first N. alessandri, a species of very limited distribution (Clarke, 1964) around 35° S. The present author (McQueen, 1976) in Table 1 ascribed this species on published evidence to the Nothofagus ‘brassi’ pollen group, but later work has shown that N. alessandri belongs to the N. ‘fusca’ pollen group (letter N. T. Moar p. 266).
The second species, although apparently formerly of wide distribution, is now limited to stands near Concepcion. This is N. glauca, a species apparently similar ecologically to N. obliqua.
A third species which was not mentioned in my previous contribution, N. leoni, is again of a very limited distribution in the province of Maule (35° 40′ S) and is a deciduous tree with leaves intermediate in size between N. alessandri and N. glauca (F. Schlegel S. pers. comm.).
Visits to these forests accompanied a project on N. antarctica generously financed by Victoria University of Wellington, the N.Z. University Grants Committee and by Le Ministère des Affaires Etrangères, Paris. I also wish to express my gratitude to Dr. F. Schlegel S., Director of the Instituto de Silivicultura y Reafforestacion
Mr. Olaf John, of Wellington, reproduced from colour slides all photographs in both Parts I and II of this series and merits my particular thanks.
In the Southern Andean region there are very distinct forest formations: Reichle (1907), Skottsberg (1910, 1916), Schmithusen (1956), Oberdorfer (1960), Hueck (1966). The diverse forest pattern can be related only partially to soil. Apparently the ecology of these forests is differentiated predominantly by climatic factors. Variability is strongly marked in the forests of South Chile between 37° and 43° S, where forests of Nothofagus are physiogonomically important. Certain species of Nothofagus are much in demand for timber from the natural forests, and Nothofagus may play a very important role in the future if afforestation with native species is begun.
Species of Nothofagus are spread over nearly the whole spectrum of forest associations and its species are, to a certain extent, mutually exclusive. Thus they can be considered differential or characteristic species for particular sites. Starting with this hypothesis an examination has been made of ecological differentiation in relation to temperature of Nothofagus. The final composition of a mature stand of trees has had a life history passing through a youthful phase of selection which is important when reasons are sought for the presence or absence of a species. Measurements were, therefore, carried out only at the level of the lower vegetation, so this research is limited to the thermal requirements and tolerances of Nothofagus saplings and seedlings.
Traditionally the climate of South Chile is regarded as oceanic and strongly humid. In an ecological sense this can be accepted only with considerable restrictions (Schwabe 1956, Weischet 1970, Weinberger, Romero and Oliva 1972). Recently more differentiated climatic classifications were proposed. Van Husen (1967) distinguishes two zones within the research area of this paper. Between 45° S and 41° S summer rainfall occurs regularly even in the lowlands. This zone extends to the north as a tongue through the mountains (Fig. 5). Between 41° S and 38° S in the Central Valley is the zone of occasional summer drought. Here it is only during the winter that the westerly wind is dominant, while summer weather is more often locally determined. Van Husen (1967) arrives at this conclusion on purely climatic criteria; this conclusion can be confirmed by plant ecology, for field observation interprets the forest diversity as a result of the varied topography of the region. Its very broken relief causes a
Temperature variations are exceptionally strong under an open sky. Because of the latitude there is intense solar radiation with consequent high temperatures in open sites during daylight but considerable heat loss by radiation at night. During summer, minimum temperatures below freezing point are recorded even at a short distance from the coast. Such a condition must be given special importance, implying a very strong selection pressure. These extreme temperature conditions occur mainly during the summer. This is shown by the annual curves of some parameters that characterise the heat and radiation relationship of a sea coast station (Fig. 1). Inland, especially in the rain shadow of the Coastal Range, the summer is even drier and warmer (Fig. 5).
In this connection the duration of sunshine and total radiation are important. The magnitudes of both influence soil temperatures considerably, and also affect the climate of the air layer close to the soil. In Valdivia on clear summer days solar radiation of up to 400 cal/cm2 are recorded. However, the minimum on rainy days in winter shows only 0.5% of this figure.
As an additional source of information comparison of the annual curves for temperature and intensity of radiation are useful. Heating at ground level is slower than heating of the air by the sun and thus corresponds to general observations. This slowing, however, is not as great as would be expected for an oceanic climate. According to Lundegardh (1957, p. 193) the temperature maximum in an oceanic climate appears only 1 1/2 to 2 months after the highest position of the sun. From this point of view there is a noticeable diminution of oceanic influence in the Chilean case.
Sites, selected for each species, were distributed in the region between 37° 30′ and 42° 30′ S, and were chosen irrespective of whether the forests were closed on open, primary or secondary. The sole criterion for the adoption of a recording site was the existence of spontaneous natural regeneration of Nothofagus.
Temperature and air humidity were recorded for periods of several days (5 to 8 days). All measurements were made in the three warmest months (December, January and February). The measurement periods for each species were distributed during different parts of the summer.
The equipment consisted of a series of ordinary thermohygrographs. They were installed on small benches in such a way that the measuring elements of the instruments were 25cm above the soil. They were calibrated against a wet and dry bulb thermometer and for additional control a minimum thermometer was installed beside each thermo-hygrograph. Because of topographic difficulty the normal type of meteorological screen for protection against direct radiation was not installed. Instead portable protection against radiation was made out of a folded sheet of galvanised iron, shaped into a tunnel 50 X 50 X 50 cm.
It was installed with the openings on a north-south orientation so that even a low angle of sun was excluded from the shade. Daily extremes and 2-hourly values for temperature and relative air humidity
The average daily temperature: expressed as the mean of a total of 12 readings (0 to 24 hours); like the next value it typifies the heat requirements. By this measurement were obtained indications of the optima and ecological ranges of a species in north to south, and altitudinal distribution.
Mean temperature of the warmer half-day (10 a.m. - 8 p.m.): In connection with other questions the water budget, and stress were of particular interest. Thus the mean temperature of the warmer half-day was determined as well. The comparison of a large number of daily curves showed that air temperature at 10.00 a.m. is generally lower than in the evening at 8 p.m. (Fig. 4). For this reason a calculation was made of the average for the warmer half-day from six temperature readings between 10.00 a.m. and 8.00 p.m. The apparent daily retarding of heating effects is of an artificial nature and due to Chilean Summer Time.
Average saturation deficit of the warmer half-day: Atmospheric saturation deficit can be used to characterise evaporation conditions. Temperature, relative humidity and saturation deficit of the air are dependent on each other. With the recorded data it was possible to read corresponding saturation deficit from tables in mm of water vapour tension. Average saturation deficit is a better characteristic than relative atmospheric humidity for defining the evapo-transpiration stress in the water budget of plants (Stocker 1956). The six values, obtained from 10 a.m. till 8 p.m., were also converted into a mean value.
Average daily temperature variation: The daily amplitude is suitable to identify the ‘continentality’ of the site. This figure was obtained from the equation:
Average daily variation = 2Ma — Mi' — Mi"/2
(Ma = daily maximum, Mi' = minimum of the preceding night, Mi" = minimum of the following night.)
Average species-specific day curve (Fig. 4): All biotopes in which a species maintains itself show similar temperature climatic features. For this reason the measurements from all sites possessing regeneration of a particular species have been combined and evaluated together. Thus the idea was developed of characterising the ‘ideal site’ with the aid of this species-specific data collection. All corresponding daily curves have been summarised in this way. The results should be interpreted as average values that identify the optimal conditions for natural regeneration. This is equally valid for general heat requirements (temperature summation) as for the corresponding daily amplitudes of temperature.
Frequency distribution of minimum temperature values (Table 3): The presentation alone of mean values results in considerable loss of information. In the present case it was valid as a rule that the measurement periods from at least 7 stations were available for each species. Thus it was possible to evaluate the corresponding dispersion of values. The probability with which night temperatures during the warm period of the year drop below defined low limit thresholds was calculated for each ‘ideal site’.
The experiment assumed that the dispersion was subject to a normal distribution. To test this hypothesis a larger data collection was needed but these data were not available for the minima at 25 cm above ground at any one site. However, for a series of years the lowest temperature at 5cm above soil was measured for Valdivia. (A. Huber pers. comm.). A close functional relationship exists between both magnitudes of parameters. Fig. 2 shows the frequency distribution for these values for 33 summer months. The total number of 993 daily values can be accepted as a representative random sample. The total data collection shows a close approximation to the normal distribution. The top of the curve indicates that apparently a mixture of at least two individual dispersions has been plotted. The study of the reason for this anomaly is a climatic problem not further examined here. It is, however, worth mentioning here the possibility that the assumed individual ranges correspond to different types of extreme weather conditions.
In this connection interest is concentrated on the area of fewer observations on the left sector of the curve, where there is a noticeable deviation from normality. The question arises if this deviation is too serious. Therefore with the aid of the given parameter, the theoretical expected frequency for the occurrence of low temperatures was calculated and compared with the empirical values (Table 1). It must be recalled that by the establishment of species-dependent total collection, an abstract of real sites is accomplished. Each of the samples of data that are combined to give species-specific collections could be integrated from different weather condition-dependent ranges, as is expected for the Valdivia station (Fig. 2). The different sites of any species, however, deviate a little from average values, being partially somewhat warmer or cooler. Consequently the distribution curves for individual sites are shifted in relation to each other. This circumstance leads to a more balanced picture for the ‘ideal site’ after combination of data and thus the results can be accepted as satisfactory.
Temperature gradients close to the soil are very steep (Geiger 1961). The data for the Valdivia station show the relationship represented in Table 2, showing the advisability also of considering temperatures just above freezing point.
Preparation of ecoclimagrams (Figs. 6 and 7): Dispersion of individual data around the mean values has also been considered for the determination of tolerance ranges peculiar to the various species. Derivation of the latter is demonstrated in Fig. 3.
Every point within the system of co-ordinates corresponds to a pair of individual values. The collection of data belonging to any species results in a distribution of points whose extension still cannot be taken as an expression of the mean ecological range. In the case of Nothofagus obliqua the lowest average daily temperature observed was 8.3° C; for N. pumilio the highest was 11.8° C. These values represent extremes which occur exceptionally during the summer in the corresponding biotopes only. The mean ecological ranges of both N. obliqua and N. pumilio are more restricted and it is for this reason that they are never observed together.
A statistical analysis led to the conclusion that about half of the total dispersion is caused by the normal variability of climatic parameters observed in any given locality. Only the remainder corresponds to local differences. In other words, only one half of total value dispersion characterises the capacity of species to adapt themselves to different biotopes.
In accordance with this assumption the average ecological ranges of species (Fig. 3, broken lines) are obtained by reducing total value dispersions (unbroken lines) to half. The reference points are given by the intersection of corresponding mean value co-ordinates. The resulting graphs are designated as ecoclima-grams.
The deciduous ‘roble’ is distributed on the Argentinian side of the mountains only between 36° 50′ and 40° 15′ S (Hueck 1966). In Chile this species advances considerably further north, in its var. macrocarpa to approximately 33° S (Munoz 1968). Here it occurs between 750 and 2,200 m altitude. In the south, dow nto 41° 20′ S the main distribution of this species is in the lowlands.
Fig. 5 shows a schematic transect across the Coast Range near Valdivia. This is the only known region where all species of Nothofagus are together with the exception of N. pumilio. Here Nothofagus obliqua goes up to about 500m on the lower rainfall east face. Generally it can be considered as the dominant species of a tree association which is typical for a region of episodic summer drought. Frequent evergreen companions are Persea lingue Nees and Laurelia sempervirens Tul.
Fig. 4 shows that Nothofagus obliqua is a relatively thermophilic tree. In its ‘ideal biotopes’ the day and night temperatures were higher than in those of all other Nothofagus species. Accordingly N. obliqua is found further into the sclerophyllous woodland where conditions are close to the mediterranean climate areas of California and the Mediterranean. The ecoclimogram in Fig. 6 reflects the tolerance range of this species in respect to north-south (vertical axis) and west-east distribution (horizontal axis). These compass orientations are obviously not to be interpreted in a strictly literal sense for it is not only the geographic co-ordinates which determine the degree of increasing latitude or ‘southerness’ and continentality of a site. These orientations have been transferred to an ecological sense, because localised topography is a deciding factor. In this way flat land will show a stronger continentality than adjacent slopes; the degree of southerness of a slope will be determined by its aspect. Fig. 6 shows a certain continental tendency exhibited by Nothofagus obliqua as does Fig. 4, with a wide temperature amplitude. It has been observed that Nothofagus obliqua never goes down to the west coast, although in breaks in the Coastal Range, near Valdivia, and at Rio Tolten the species grows within a few kilometres of the sea. This species does not penetrate into the coastal belt of Aextoxicon forest. Even in open pioneer communities this light demanding species is absent near the sea. Nor does it grow on the flat foreland of Arauco that extends in the front of the coastal range between 37° 15′ and 38° 15′ S.
In the other ecoclimogram (Fig. 7) there are two parameters (temperature and saturation deficit) that are linked by a positive correlation. The higher are their values then the drier and warmer are the respective sites in summer. It must be recalled that these
Nothofagus obliqua is considered an indicator of fertile sites in the southern part of its distribution, but in the north, N. obliqua occupies sites of shallower soils. Because there are commonly long summer droughts in the hardwood region N. obliqua has to be well adapted to this contingency. In this respect, Fig. 7 shows a certain similarity between N. obliqua and N. antarctica.
The ideal site for N. obliqua, even though relatively warm is not, however, free of frost (Table 3). For south Chilean agriculture this is an important factor because former sites of N. obliqua forest are now intensively farmed. Here summer frosts quite often cause serious damage to wheat during the flowering season.
N. obliqua prefers flat sites but not to the same extent as the next species, N. antarctica. This is true not only for the flat lands of the Central valley. For example one can find N. obliqua in the northern sector of the Coastal Range (Nahuelbuta National Park, 37° 47′ S 73° W) on high plateaus of 1,200m altitude. There its range overlaps with that of Nothofagus antarctica and Araucaria araucana C. Koch.
N. obliqua does not grow on the slopes of the Coastal Range; instead one finds N. procera (syn. N. alpina) with numerous evergreen species. In the lowlands, 1,000 m below, Nothofagus obliqua is again found and has the same type of distribution in the Tolhuaca National Park (38° 12′S 71° 48′W) in the Andes.
By contrast to Nothofagus obliqua the nirre, which is likewise deciduous, is a slow growing tree or shrub seldom over 12m high. Its distribution in Chile is from 35° 31′S and in Argentina from 38° S, thence southwards to Tierra del Fuego (Urban 1934, Skottsberg 1916). The ‘ideal’ N. antarctica stand is characterised by particularly high daily temperature variation (Fig. 4). On one hand are day temperatures less than 2° below the peak of the thermophilic
N. antarctica is also observed in the southern part of the area of distribution of N. obliqua. Only occasionally are both species found together, there being generally a sharp boundary between the two forest types, so that the transition from one to the other may take place within a few paces. In such conditions the areas occupied by N. antarctica commonly are flat valleys with shallow soils, areas called locally ‘nadis’ (Weinberger et al. 1970). Such sites are occupied by ‘Zarzales’, formations which are open and contain a high proportion of thorny shrubs, Berberis, Discaria etc. Embothrium coccineum is frequently present and shows a preference for such sites along with Nothofagus antarctica.
N. antarctica is a notable light demander and in this respect is very similar to N. obliqua. This behaviour is reflected in the high values for both species of the average temperature of the warmest half-day (Fig. 7). Sites studied are frequently open and subject to strong insolation. However, both species show quite different optima and tolerance ranges. This is expressed not only by the data in Table 3, but also in Figs. 6 and 7. This is especially the case for the ranges of high temperatures and saturation deficits; there the ecoclimagrams demonstrate noticeable divergences between the two species. The temperature environment which corrsponds to the cuneiform intercalations between the ranges of the two species (Figs.
Austrocedrus chilensis and Lomatia hirsuta.
N. antarctica sites are characterised by particularly high atmospheric saturation deficits. There the soils are usually only partially covered by vegetation. They dry out on the surface very rapidly
N. antarctica is the only species of the genus which tolerates conditions of the sub-Andean rain shadow steppe formation.
The ‘rauli’ forms almost pure stands as trees up to 40 m in height. It is distributed in mountain areas of Argentina (39° 22′ to 40° 23′S, Hueck 1966), and Chilen (35° 20′ to 40° 20′S). This
Nothofagus species.
According to Figs. 4 and 6 the continentality is approximately the same as for N. obliqua. Mixed stands are located in areas of transition between the two species but the ‘ideal’ stand of N. procera is considerably cooler than for N. obliqua. In 46 samples from stands of N. procera, N. obliqua was present in less than one third. A marked plant-sociological link exists with N. dombeyi which was present in two-thirds of the stands sampled. Coincidentally in Fig. 4 is shown a large similarity of the temperature curves of both species (N. procera and N. dombeyi).
N. procera frequently occurs with Lomatia dentata. In general the deciduous N. procera shows considerable affinity to various evergreen trees of the cool oceanic climate. This affinity goes so far that in Valdivia province, on the eastern flanks of the Coastal Range at 800-900 m this species forms mixed species — rich stands with N. nitida (Fig. 5). According to Urban (1934) N. procera appears in the province of Chiloe scattered in the evergreen forests. However, the writer did not succeed in relocating any N. procera there.
According to the ecoclomagrams, the thermal requirements of the sites of N. antarctica and N. procera overlap to a considerable extent. Nevertheless both species are not found growing together because there are other differentiating factors. N. procera is almost exclusively a tree of slope positions, on soils developed in the Andes from recent volcanic ashes. On the geologically older Coastal Ranges it is limited to deeply weathered soils, where competition pressure on these soils is very strong. The excellent ability of N. procera to succeed under such conditions is shown by the fact that it shows good regeneration in heavy shade. By contrast, it has already been emphasised that N. antarctica is light demanding.
N. procera is a species with large and relatively soft leaves, so that more than any other Patagonian species of Nothofagus it resembles northern hemisphere summer green trees. It is dependent on a considerable soil water reserve being available and would be unable to survive on N. antarctica sites.
Finally, the frequency of frosts is substantially lower in N. procera sites and is a consequence of the temperature regime of slopes occupied by this tree. The interpretation of many daily curves of temperature show that on these sites at night warm air streams flow upwards from the valleys (Fig. 4). Such air flows contribute to a marked lowering of the night cooling effect and may often result in temperature increase. This phenomenon is so marked that it remained even when the average from 62 daily temperature curves was calculated. Undoubtedly this is a mechanism which helps to reduce the number of frosts. This phenomenon is responsible for the fact that the relative humidity of the air of the ‘ideal’ sites for Nothofagus procera is approximately 90% between 4 a.m. and 8 a.m. whereas for all other Chilean Nothofagus, including Nothofagus obliqua, there is a variation in the corresponding mean values, from 95% to almost 100% relative humidity. Consequently dew is scarcer on sites of N. procera.
The occurrence in the same areas of N. procera and N. pumilio is not known to the writer. According to Fig. 6 this would not be expected.
The ‘lenga’, a summer green species reaching 30 m height, forms typical tropophilic forests, according to Schimper's definition. These forests are sharply delimited from all other woodland associations plant-sociologically, and extend with particular uniformity from 35° 30′S in Chile and 36° 56′S in Argentina to the southern end of the continent. In Tierra del Fuego this species grows at sea level (Skottsberg 1916).
N. pumilio and N. antarctica are often mentioned together because of their similar Andean and Subantarctic distribution. However, the ecological requirements of these species are very different as is shown, primarily, by the daily temperature curves (Fig. 4). Where N. pumilio regenerates well no high temperatures are recorded in the midday hours. Regeneration is essentially on shaded sites, where the saturation
Fig. 4 also shows that the mean night temperatures for both N. pumilio and N. antarctica are very similar. Nevertheless even here there is a characteristic difference in the smaller variability of the temperature minima for N. pumilio sites. Consequently there is a significantly lower frequency of night frost (Table 3). From the fact that the average daily temperature for N. pumilio is generally within a lower range than that of N. antarctica (Fig. 6) it follows that the latter possesses greater plasticity. It is probable that the restriction of N. pumilio in the northern part of its area to the more moderate microclimates of slope sites, is due to the thermal limitation described above.
In this connection it is of interest to compare leaf emergence, which in Nothofagus pumilio occurs up to two weeks earlier than for N. antarctica. N. pumilio flowers and begins to open its leaves when N. antarctica is still leafless. This phenological difference, which was noted by Skottsberg (1916) is generally very marked and also indicates a better adaptation by N. antarctica.
In the southern part of its area N. pumilio also occupies valley flats and formed extensive forests; around Punta Arenas these forests are mainly destroyed by farming activities. This distribution thus raises the question of how this biotope change can be understood. Data on Table 4 and comparison with more northern stations support the following explanation:
Lonquimay at ca.40° S is in an Andean high valley in which the extensive N. antarctica low forest is physiognomically important. N. pumilio is located here only on the higher slopes. The sky is usually cloudless during the warmer part of the year. By contrast the sky of Punta Arenas is overcast at least every third day. Such cloudiness, significantly frequent at Punta Arenas, demonstrates weather conditions which prevent a strong inward or outward radiation. As a result there are considerably smaller daily temperature variations at Punta Arenas.
Moreover, northern sites of N. antarctica owe their thermal characteristics to the circumstance that they are excluded from the equalising influence of the west wind. In the south the effects of such winds are more pronounced, even inland, and for that reason there are extremely pronounced allochthonous climates (Weitschet 1970). The low number of calm days in Punta Arenas, by comparison with Lonquimay, shows that there is a permanent intensive exchange of air masses at Punta Arenas, and the formation of static cold air ponds is less probable. As a result of these conditions there are practically never any night frosts detected during the warm period of the year at Punta Arenas (at 2 m above soil). However, 1700 km to the north, at Lonquimay such frosts on an average occur every tenth summer night.
Nothofagus pumilio forms the timber line on the Andes more frequently than other species. Above is a dwarf scrub, very rich in Ericaceae. The plant-sociological link of N. pumilio forests to the evergreen forest is slight, but of special importance are Drimys winteri, Maytenus magellanica, Nothofagus betuloides in southern areas and Nothofagus dombeyi that comes from lower altitudes into the Nothofagus pumilio forests in the Andes.
The evergreen ‘coigue’ is distributed in Chile from 35° S to 47° 30′S (Skottsberg 1916) and in Argentina from 38° to 44° S (Dimitri et al. 1950). According to Fig. 4 and 6 it is a tree with its main distribution on sites of oceanic tendency. Growing to a height of up to 45 m, it reaches its optimum development in the high rainfall western slopes of the Andes mountains.
According to Hueck (1966 p. 364) it is a demanding species that needs deep and dominantly moist soils. The present author wishes to modify this opinion. In fact its ecological amplitude is considerably wider, as Nothofagus dombeyi penetrates successfully into the biotopes of all the other species considered in this paper. In the area examined no other species shows a comparable adaptability. In the region of Valdivia, with increasing altitude or on shallow soils of the lowlands there is an impoverishment of the species-rich mixed forest which gives way to almost pure N. dombeyi forest. Its relative insensitivity to cold goes so far that this species goes up to the lower part of the winter snow line zone. For example, in the National Park of Puyuehue at 900 m to 1000 m there are extensive mixed forest with N. pumilio. In the lowlands N. dombeyi penetrates into the areas of frost exposed ‘nadis’ where it is established together with N. antarctica.
The volcanoes Llaima (38° 42′S), Villarrica (39° 23′S) and Osorno (41° 06′S) lie west of the main range of the Andes in a more oceanic climate. N. dombeyi is located here as a pioneer species on lahars (the volcanic debris streams which flowed down from the volcanoes after eruptions had thawed the ice). Lahars represent extreme sites where soil formation is at an initial state. Most species are not able to establish except with the protection of scattered N. dombeyi shrubs. Such sites are essentially free of vegetation and exposed to strong insolation. Therefore high water saturation deficits are frequently observed in the air layer close to the soil. These conditions, however, usually are limited to short daily periods because such localities are open to west winds, normally starting at noon and bringing moist air masses from the Pacific Ocean.
Less frequent in South Chile are weather conditions defined by a warm dry wind locally called ‘puelche’, occurring every summer for some time. This wind advances over the Andes from the eastern grasslands and causes, on open mountain sites, an abrupt temperature
The La Picada site (Fig. 8) is one of such lahars. This site is on an open mountain saddle. The fact that N. dombeyi, usually considered as a ‘rain forest species’, regenerates well at this site shows that it is exceptionally well adapted. In fact N. dombeyi, in a comparison of 53 evergreen species, had the highest plasmatic drought resistance (Weinberger et al. 1972).
Taking into account the above mentioned results, N. dombeyi can be characterised as follows:
It is a tree with lower tolerance limits to xerothermic conditions than the summer green N. obliqua, N. antarctica and N. procera (Fig. 7). However, N. dombeyi resists considerable atmospheric and physiological saturation deficits, if they are transitory and when the overall climate of the site is oceanic. Within the more continental N. procera belts of the mountains it is generally observed that N. procera prefers the exposed slopes whereas N. dombeyi is found in the wetter more temperate mountain gorges (Fig. 5).
This evergreen tree closely resembles the preceding species, N. dombeyi and, its common name is ‘Coigue’ or ‘Coigue de Chiloe’. Its distribution starts from 40° 20′S on the Coastal Range near Valdivia (Fig. 5) southwards to 48° 30′S (Skottsberg 1916). It has not been recorded on the east of the Andes.
Although the daily temperature curves of N. dombeyi and N. nitida are similar (Fig. 4) the climatic limits of N. nitida are
N. dombeyi (Figs. 6 and 7). However, it is only occasionally that both species are found in mixed stands as they differ very markedly in their edaphic requirements. N. dombeyi grows well on the wet soils of depressions in the northern part of its area, whereas towards the south a biotope change is observed. On Chiloe Island the wet soils are occupied by extensive N. nitida forests, whilst N. dombeyi is limited to the better drained morainic soils. Further to the south N. nitida is the tree of the flat and permanently wet soils along the coast.
In further comparison of both species the wider tolerance range of N. dombeyi to daily temperature variations (Fig. 6) is essentially based on its adaptation to higher temperature maxima. The dispersion downwards of minimum values is only a little higher for N. dombeyi sites. Finally the average night temperatures for N. nitida are a little below the temperatures for the other species (Fig. 4). Thus there is only a slight difference in the amount of summer night frost (Table 3). As mentioned earlier, even moorland sites in the coastal area can be affected by a certain frost frequency. This applies to West Patagonia where N. nitida occupies flat coastal lands.
The biotopes of N. nitida are defined by permanently high air humidity. Accordingly this tree occupies the high rainfall belts of the montane Cordillera Peladar (part of the Coastal Range) (Fig. 5). In this respect N. nitida is more demanding than all other species considered (Fig. 7), and may be related to the fact that this species has a very shallow root system.
N. nitida shows strong phyto-sociological links with other forests of evergreens, particularly with Drimys winteri, Weinmannia trichosperma, Laurelia philippiana, Amomomyrtus luma and the conifer Saxegothea conspicua. N. nitida is better suited to very temperate climatic sites than N. dombeyi and is present adjacent to one of the few forest formations of south Chile where the forests lack any Nothofagus species. Such are the forests of the western flank of Coastal Range which are characterised by many trees of the Myrtaceae. Southwards of Valdivia such forests increasingly replace Aextoxicon forests (Fig. 5). However, such Nothofagus-absent forests do not penetrate the general area of the following species.
The evergreen ‘Ouchpaya’ or ‘Cohue de Magallanes’ dominates coastal areas from 48° southwards. In the Valdivia region it is found only in a few isolated stands located on the highest and most windy position on the Coastal Range (Fig. 5), and inland, on the volcanoes Osorno and Calbuco. This tree does not grow tall and in the main forests of the Magellanic areas few trees are taller than 15 m.
In south Patagonia Nothofagus betuloides forms the ‘maritime forest boundary’ inland of the oceanic heath and moorland (Skottsberg 1916). Table 4 defines the climatic conditions. Evangelistas is one island on the outer coast where N. betuloides can grow only in the protected hollows. Temperature variations are very small and clear days are scarce on these coasts where there are storm winds at all times of the year. Even in summer wind speeds of over 40 km per hour occur every two days on the average. N. betuloides is well adapted to such an environment, with its compact growth and particularly small leaves.
Unfortunately there are only 11 daily measurements for 2 sites and for this reason it is only possible to estimate its ecological tolerance. Only the average values which could well define the ‘ideal site’ are represented in the ecoclimagrams (Figs. 6 and 7). The position of N. betuloides in these diagrams demonstrates that it is the equivalent in the far south to the two other evergreen species (N. nitida and N. dombeyi). On the other hand it is in the whole of its distribution in contact with stands of N. pumilio which grows on higher sites on mountains and drier sites to the eastward.
In its habit N. betuloides is very similar to stunted examples of N. dombeyi. The climatic tolerance appears similarly wide and certainly greater than that of N. nitida. However, N. betuloides does not grow on such dry sites as Nothofagus dombeyi. In the trans-Andean valley of the Rio Baker (in Aysen Province, approximately 47° 50′ S) N. nitida occurs on the coast; somewhat further inland is N. betuloides, and 60 to 70 km inland from the mouth of the river a zone of N. dombeyi commences. Skottsberg cited this succession as very remarkable. ‘It is surprising to find Nothofagus betuloides replaced by Nothofagus dombeyi inland when the opposite would be expected’ (Skottsberg 1916, p. 71).
However, according to the results of the present study the plant geographic observations of Skottsberg are quite comprehensible. It had already been noted that a warm and very dry east wind (Fig. 8) can penetrate such openings in the mountains as the Rio Baker. The average temperatures between 10 a.m. and 8 p.m. (Fig. 7) indicate that N. betuloides does not tolerate high temperture to the same extent as does N. dombeyi. The distribution in the Cordillera Peladar, where both species are represented, also shows clearly that Nothofagus dombeyi tends towards an inland climate, rather than Nothofagus betuloides.
However, in even more southern and colder regions N. betuloides substitutes completely for the other species. It is there established on edaphically favourable sites as well as to the east of the Andes (Hueck 1966).
Dear Sir,
In Table 1 of ‘The Ecology of Nothofagus and Associated Vegetation in South America, (Tuatara 22: 38-68) Nothofagus alessandri to the ‘brassi’ group.
Pollen of the rare N. alessandri is usually placed in the fusca group (Cranwell 1939) but van Steenis (1971. p. 97) raises a doubt when he comments that N. alessandri pollen should be referred to the ‘brassi’ group. an opinion based on a private communication from Dr Archangelsky, Museo de Ciencias Naturales, La Plata, Argentina. Heusser (1971, p. 35, pl. 28-334) describes and illustrates pollen of a specimen, SG 63329, Museo Nacional de Historia Natural, Santiago, identified as N. alessandri which is clearly of the ‘fusca’ type. In a letter (5 April. 1976) in response to a query from me, Heusser states that he has ‘no reason to suspect that N. alessandri represents “brassi” type’ and that therefore his findings are ‘in agreement with Lucy Cranwell's interpretation’. Cranwell (1939) referred pollen of N. alessandri to the ‘fusca’ type, and a duplicate slide of her material in the Botany Division collection supports that conclusion. Van Steenis believes that Cranwell's material, obtained by Skottsberg from a specimen in the herbarium of Arnold Arboretum, was wrongly identified, but this seems unlikely in view of Heusser's (1971) more recent work with a different specimen.
In the present state of our knowledge then it seems that N. alessandri pollen should be assigned to the ‘fusca’ group and that ‘brassi’ type pollen is not now represented in South America, although there is no doubt of its presence in the Tertiary (Cookson and Cranwell 1967). It is important to draw attention to this matter since there is considerable interest attached to the biogeography of Nothofagus in the Southern Hemisphere.