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Tuatara aims to stimulate and widen interest in the natural sciences in New Zealand, by publishing articles which (a), review recent advances of broad interest; or (b), give clear, illustrated, and readily understood keys to the identification of New Zealand plants and animals; or (c), relate New Zealand biological problems to a broader Pacific or Southern Hemisphere context. Authors are asked to explain any special terminology required by their topic. Address for contributions: Editor of Tuatara, c/o Victoria University of Wellington, Box 196, Wellington, New Zealand. Enquiries about subscriptions should be sent to: Business Manager of Tuatara, c/o Victoria University of Wellington, Box 196, Wellington, New Zealand.
James Adams was born near Killarney, Co. Kerry, on May 12, 1839, and was the eldest son of Alexander and Edith Adams. In common with the whole of the Irish countryside the family was deeply affected by the potato famine that resulted in great hardship for the people on their land and an almost total loss of income for the landowner. James was educated in Ireland and as a young man went to London where he took a B.A. degree at London University. In 1865 he obtained the post of headmaster at a privately endowed grammar school at Douglas on the Isle of Man. Although well versed in Latin and Greek from an early age he was against boys spending so much time on the classics when chemistry, physics, mathematics and, of course, natural history were, to his mind, very important. His forward-thinking ideas on education, particularly that it should be free, secular and for girls as well as boys, led him to give up a pleasant and comfortable life to emigrate to New Zealand. With his wife, Ann, and four small children he left Liverpool in September, 1870, on the steamer ‘Great Britain’ for Melbourne. They intended to settle in Dunedin where the educational system of the Scottish settlement seemed to be very close to his ideals. The steamer ‘Gothenburg’ brought them to Port Chalmers but to his disappointment he was unable to secure a teaching post and was forced to use his letters of introduction to the Bishop of Auckland. Within a short time he and the family were living in Parnell following his appointment as assistant master at the Church of England Grammar School at St. Mary's Cathedral. In 1872 he succeeded the Rev. Dr. Kinder as headmaster and remained so for the next eight years. It was at this time that he formed a long and close friendship with
Immediately the rugged bush-clad hills of the Coromandel Peninsula claimed his interest and he followed Thomas Kirk's account of the botany of the Thames goldfields with his own observations published in the Transactions of the New Zealand Institute in 1883. During the years 1881, 1882 and 1883, often with his eldest son, Ernest, then a surveying cadet under Percy Smith in Auckland, he climbed many of the high peaks on the peninsula — Table Mountain (Whakairi), Kaitarakihi and Maumaupuki amongst them — and crossed the peninsula to Mercury Bay and Tairua. Two notable plant finds were Celmisia adamsii from the crags of Table Mountain and Castle Rock and Elytranthe adamsii from the Hape Creek above the township of Thames (then consisting of two settlements, Shortland and Grahamstown, collectively called ‘the Thames’).
It seems that Cheeseman and Adams had a plan to visit all the isolated high hills in the Auckland district and together or separately they carried this out over a number of years. Pirongia in 1879, the peaks south of Coromandel between 1880 and 1883, Te Aroha, 1884, were those that Adams visited. The most significant discovery came in January, 1888, when with his son, Ernest, then surveying near Cabbage Bay, he climbed to the high peak of Te Celmisia incana, Podocarpus nivalis, Pentachondra pumila, Carpha alpina and Ourisia.
Other botanical trips were made to various parts of New Zealand — the Mt. Arthur Plateau with Cheeseman and Meyrick the entomologist in January, 1886; Mt. Hikurangi and the east coast with Petrie in January, 1897 (on this trip they were accompanied by J. Lee, a teacher from the Native School near Hicks Bay and two Maoris, Winiata and Morgan); North Cape with Cheeseman in 1895; and the Mt. Cook district and Lake Tekapo with Cheeseman in January, 1898. Herbarium specimens show that he also visited the Volcanic Plateau, Mt. Egmont and Castle Hill, Canterbury, but no field notes for these trips remain.
Podocarpus halli Kirk is named. Hall exchanged seeds of native trees with friends in Britain over many years including the Dorrien-Smith family of Tresco in Cornwall where many New Zealand plants flourish. His own arboretum of native and exotic trees remains in part amongst a new housing estate on the hillside above the south end of Thames (see Trans. N.Z. Inst. 34, p. 388).
In 1906 Manual of the New Zealand Flora for which all of his own botanical observations and collections had been made in the hope of assisting his friend. His small herbarium was presented to the Auckland Institute and Museum but because, it is said, a well-meaning daughter tidied away many scribbled notes and labels, the specimens have been left with fewer details than he would have provided. With the herbarium are some field notebooks covering some of his Thames collecting trips, and the Mt. Arthur, Mt. Hikurangi and Mt. Cook expeditions; regrettably the remainder have been lost.
The Date of publication of Thomson's well-known work is a matter of some interest in respect to a problem of zoological nomenclature. This problem of priority is somewhat complex and the taxonomic aspects have been considered elsewhere (Dawson, 1971).
In brief, the Tertiary to Recent brachiopod originally named Terebratulina cancellata Koch was renamed Terebratulina hedleyi by Cancellothyris australis. The need for renaming arose from the preoccupation of the specific name by Terebratula cancellata Eichwald, 1829. Both Finlay's and Thomson's names have been used subsequently.
Since Thomson's 1927 monograph also contains eight new generic names (Hispanirhynchia, Abyssothyris, Japanithyris, Jaffaia, Pictothyris, Neobouchardia, Pirothyris, Malleia) which have been used subsequently and might well be involved in priority questions, I have made some effort to determine the actual date of publication of Thomson's ‘Brachiopod Morphology and Genera’.
This book was published by the New Zealand Board of Science and Art, under the imprint of the Dominion Museum (
The archives of the Dominion Museum hold some correspondence and papers of Dr.
In a letter, dated May 26, 1927, to Professor Ichiro Hayasaka of Tohoku University, Thomson stated:
‘I am very shortly bringing out a manual of the Morphology and Genera of Recent and Tertiary Brachiopods, and it would be perhaps as well for you to see it before completing your revision’ [of the Tertiary Brachiopoda of Japan].
To J. Wilfrid Jackson, Thomson wrote (September 15, 1927):
‘I have delayed answering your letter until I could send you a copy of my book…. It is now being printed off by the Government Printer, but may be a little delayed because at this time ofthe year there is a pressure of Parliamentary printing at the printing office. Meanwhile I send you page proofs. I am having the greatest difficulty in getting the figures straightened up — I fear they will disfigure the book. I have arranged to be allowed to distribute a number of free copies to correspondents who will consent to review the book in some suitable scientific journal, and hope you will accept one on these conditions. You are in a better condition to judge of the value of the new book, and to appreciate what parts are original and new, than any one else. I hope its perusal will tempt you back to the group…. Now that the book is off my hands I hope to … tackle a palaeontological bulletin on the New Zealand Tertiary species. My health, however, prevents me making very quick progress, for I have just had to take another three months sick leave, and have to nurse my powers very carefully. I am sorry to see the ranks of Brachiopod scholars rapidly thinning. I had hoped Clarke and Walcott and Dall would live to see my book. Schuchert tells me he has a young man taking up the group, who will work on his very extensive collections and test out my new proposals in classifications and phylogeny, which he does not apparently like. I am not exactly old yet and am rather amused that a tyro is to put us all right …’
The ‘tyro’ was Dr. G. Arthur Cooper, now the world's leading brachiopod specialist, lately retired from the U.S. National Museum as chairman of the Department of Paleobiology (cf. Dutro, 1971); how prophetic Schuchert's words to Thomson were is well shown by Cooper's latest work, a fundamental analysis of the criteria on which generic classifications in the Brachiopoda have been established (Cooper, 1969).
On November 28, 1927, Thomson wrote to S. S. Buckman:
‘At last my book … is due from the printers, and a copy will be posted to you by an early mail. The book is issued by the Board of Science and Art, but I have secured free copies for my correspondents mainly on the understanding that they will treat them as review copies. In your case no such stipulation is made, but if you care to write a review and send it to one of the Science journals, I shall be very grateful, as there is no more competent than yourself to do so. The paper I previously submitted for the Royal Society consisted of extracts from the book, and as Dr. Butler considered it too long for the amount of new matter contained in it (which he apparently estimated mainly by the number of new names proposed), I withdrew it and awaited the publication of the book. The reading of the proofs and checking of the references occupied most of my spare time till the middle of this year. Since then I have been again on sick leave, and unable to do any further original work. Now that the book is off my hands, I hope when health permits, towrite a paper on Dr. Mortensen's Pacific collections and then to return to my Bulletin on the New Zealand Tertiary species.’
There seems to be no further conclusive evidence in Thomson's papers of the date of publication. It would seem that the book had not appeared by the end of November, 1927. The library of the Dominion Museum possesses two copies, one presented in 1958 from the library of the late Dr.
Finally, a word must be said of the worth of Thomson's book, the completion of which may be said to have cost him his life (judging by Thomson's letter to Buckman). Thomson died, a victim of tuberculosis, on May 6, 1928, at the early age of forty-six. Contemporary reviews appeared in Nature, Geological Magazine, American Journal of Science, N.Z. Journal of Science and Technology, amongst others (see Anon., 1928; C.S., 1928; G.H.U., 1928; Jackson, 1928) and included such comments as: ‘Since … 1894 … no other work has dealt so comprehensively and so lucidly with this group’ and ‘The book, indeed, is a model of what a text book in a biological group should be’ and, a word of criticism, ‘considering the somewhat high price of the book, it is regrettable that the two plates have been printed back to back, and that their reproduction is not more distinct’. In his memorial to Thomson, Oliver (1928) wrote: ‘It is not too much to say that in all zoological literature there is not a work of its type better done.’ Thomson's father, the Hon.
Thomson's work has not been surpassed even by the magnificent two-volume Section H of the ‘Treatise on Invertebrate Paleontology’ published in 1965 but, rather, is complemented by it. Nor does Rudwick's (1970) recent book of brachiopod structure and function render it redundant. It remains a fine memorial to a great scientist
I am much indebted to the Director of the Dominion Museum for allowing me to quote the letters of Dr.
In a Continuing Study of the geology near Waikanae 14C dates have contributed to the geochronology of the last 35,000 years, when climate, vegetation, and sea-level fluctuated dramatically in response to the last two discernable cold stadials of the Last Glaciation (Otiran) and to post-glacial warming.
The time-span covered by 14C is small compared with the total spectrum of geological history, but disproportionately significant because of the importance of late events to the understanding of present geography and vegetation. What can be dated depends more on the limited occurrence of datable material (lignite, peat, wood fragments) than on the sediments one wants to date; many sequences spanning parts of the 14C range remain undatable because they contain no carbon.
The early temptation to submit samples on the chance of them being datable resulted in dates beyond the limit of the method. With more knowledge of the geochronology, later samples have all produced positive dates. In an unknown area, it is most profitable to work back in a stratigraphic framework, from known post-glacial samples towards Late Glacial, awaiting results from younger samples before asking for older ones, but this assumes a leisurely timetable for the project. Palynological examination provides so much more significant history to date than a mere lithological sequence that it is perhaps unjustified to submit 14C samples from a carbonaceous sequence unless pollen analysis has already determined its vegetational and climatic history.
The geological interest in testing the contemporaneity of coolings in different parts of New Zealand and the biological interest in vegetation history during the Otiran (‘Glacial’) Stage gives dates in the range 20 to 40 thousand years a particular importance so that improvement of the reliability of the method in this range is amply justified.
The area called the Golden Coast by land-agents is the coastal lowland between
The total span of time represented by the deposits of the district covers about 200 million years, back to the Triassic greywacke suite forming the Tararua Range and its foothills; 14C dating covers at most 40,000 years. But because we look back to the past through ‘perspective spectacles’, the period to which 14C can contribute is far more important to the geologist than any 40,000-year period in the last 199 m. years.
We can skip over the deposition of the Triassic greywackes (Fig. 1 I), their deformation and elevation in the Rangitata Orogeny (L. Cretaceous), subsequent peneplanation and transgression by Oligocene seas (40 m. years ago), and the deformation that resulted in a narrow infaulted strip of Oligocene greensand behind Otaihanga (Fig. 1, II). Later in the Tertiary, about 10 m. years ago, the deformation of the Kaikoura Orogeny began to raise the Tararua Range as a complex of high blocks (and Kapiti is a smaller one); these anticlinals were progressively elevated in relation to the synclinal or relatively subsiding area of Cook Strait and the narrower synclinal between Kapiti and the mainland.
The Quaternary Era is first represented by a formation of ancient coarse gravels (Reikorangi Gravels, III in Fig. 1) that are strongly deformed and weathered. They probably represent the early Pleistocene Wanganui Series and are in part equivalent to the Kaitoke Gravels of Hutt, the Moutere Gravels of Nelson, and the marine Wanganui Series of the Palmerston-Wanganui Basin to the north-west. At some later Quaternary date the land surface was deeply weathered
Before the later ice-ages, the land gained more or less its present outline, although tilting and faulting movements have continued. Interpretation of younger Pleistocene deposits is within the framework of several theoretical generalisations, some of which are still being tested:
Climatic fluctuations (and the deposits they controlled) were contemporaneous, in a general way, through New Zealand if not throughout the world.
During each glacial period, when glaciers advanced in the South Island mountains, the Tararua Range suffered a periglacial climate even if it did not support glaciers. Mountain vegetation was reduced to sub-alpine grassland and herb field, solifluction was widespread, and streams and rivers, overloaded with waste, aggraded their beds to a steep gradient with coarse ill-sorted gravels derived by even steeper fans from tributary streams. Small glaciers (VIII in Fig. 1) were certainly present in the highest Tararuas in the Last Ice Age (Adkin, 1910).
World-wide withdrawal of ocean water to form ice sheets and glaciers lowered ocean level to depths of some 120 to 200 m. below its present level. Because deep water is quite close to the Golden Coast (south of Kapiti), ice-age gravel plains plunged steeply to coastlines at no great distance from the present shores.
During interglacial periods, melting of ice raised sea levels generally to heights above present sea-level. The land was fully forested; rivers cut down in their inland courses. Gravels in rivers and on coasts tended to be better sorted.
Tectonic movements, continuing through the Pleistocene, have caused irregularities in the levels of glacial and interglacial shorelines but were not sufficiently strong to obliterate completely the cycle of relatively low levels for glacial and high levels for interglacial ages. Therefore marine deposits on land are almost certainly interglacial, coarse ill-sorted gravels probably glacial.
Sand dunes characterise stable or prograding coasts (not advancing seas), so that most dune-sand deposits, especially those containing mica derived by long-shore drift from Tertiary beds further north, probably date from a period of sea retreat
In the valleys of the Waikanae and other rivers terrace remnants of the penultimate and probably anti-penultimate glacial gravels (Fig. 1, IV) can be recognised as such by their altitude, dissection and strongly weathered loess soils that are developed on them. Remnants of the deposits of the Antipenultimate Interglacial are also suspected in several places. These have to be recognised and classified by geomorphological, pedological, and lithologic criteria; they are all beyond the range of 14C. In practice, so little carbonaceous material occurs in gravels formed by cold-climate aggradation that there has been little temptation to submit material from such deposits.
The most conspicuous formation of the coastal lowland has long been known as the Otaki Sandstone or Otaki Formation (Fig. 1, V) and was studied by Adkin, Oliver, and Cotton before the development of 14C techniques. Te Punga (1962) described a well-exposed section near Te Horo and collected a sample (N.Z. 65) dated as greater than 45,000 years from a lignite band. By the criteria listed above, the Otaki Formation is Interglacial, consisting of the marine beach gravels and sands of a transgressive high-level sea, the beach-derived micaceous dune sands that advanced as the sea retreated, and lignite deposited in swamps ponded by the dunes. It is backed by an old cliff-line (Fig. 2, Line A-A). As it is the latest interglacial deposit recognised on the coast, the Otaki is attributed to the Last Interglacial Age, which from isotope datings overseas is thought to be about 80,000 to 120,000 years old and thus beyond the range of 14C.
Overlying the Otaki Formation are a variety of deposits attributed to the last glacial stage: coarse ill-sorted river gravels in terraces, sometimes with erect pebbles indicating frost climate near the surface (
The fan gravels occasionally preserve thin lenses of carbonaceous silt, representing swampy deposits later buried by the growing fan (Fleming, 1970). One such deposit yielded a sequence of pollen samples indicating cold wet grasslands — shrubland vegetation with a temporary silver beech phase, dated (N.Z. 573) as 19,200 years B.P. (± 560 years), and thus falling in the period of the Later Kumara-2 glacial advance of Westland (Suggate, 1965) and (less precisely) of the Takapu Stadial of Wellington Peninsula (Brodie, 1957). The fans appear to slope concordantly to the main terrace of the
In two exposures near the crossing of the Waikanae River by Main Highway No. 1, Earlier Kumara-2 of Westland, and the Opunake glacial cooling of Taranaki (Grant-Taylor, 1964), dated as 30,000-38,000 B.P.
In the second exposure, 0.75 km. downstream, a relatively complete section of the Otiran Stage overlies Otaki Formation (Oturian Stage) (Fig. 3)
The Waimahoe Lignite (C) records a cool-warm fluctuation before the cooling that presumably led to the ‘Earlier Kumara-2 = Opunake’ glacial (D-E). The only 14C sample, collected by Te Punga from an uncertain horizon, is N.Z. 22 (more than 35,000 B.P.). If confident dating in the range 35,000-40,000 becomes possible the section is worth re-sampling for 14C, as it is a rare example of three cold stadials in stratigraphic sequence in a single exposure.
Twenty-seven miles north of Waikanae on broad surfaces above the aggrading riverbeds, Koputaroa Dunesand (a cold climate dunesand) accumulated over a span of time that included a climatic cooling indicated by pollen in peat (McIntyre, in Cowie, 1963) since dated as 35,000 ± 1700 years B.P. (N.Z. 522) (i.e. equivalent to Early Kumara-2) and also the Oruanui Ash (dated as about 20,000 B.P. by correlation and thus Later Kumara-2). Recognition of this ash shower, widespread throughout the North Island (Vucetich and Pullar, 1969), allows a valuable link with the history (and 14C dates) of other districts. A similar but younger cold climate dunesand (Te Whaka Dunesand) overlies the
Stratigraphic Table
During the Otiran, the sea is assumed to have been at low levels (c. 120-200 m.) and the coast to have been somewhere south of the Kapiti Strait synclinal, which was a deep depression receiving the gravels from Waikanae district.
As the sea level rose it flooded the Kapiti Strait depression which later was partly filled by long-shore drift and prograding deposits.
At Foxton, sea level had risen to −50m. by 9900 (± 150) years B.P. (N.Z. 81). Thick beach sands with shells (e.g. at −63m. at Paraparaumu Aerodrome) presumably represent this period of rising sea level, but have not been dated. They are represented by IX on Fig. 1, and have been named Kenakena Formation. Post-glacial sea level rose to its maximum at about 5140 ± 90 years B.P., a date (N.Z. 519) from the beach deposits (Paripari Formation) near Centennial Inn,
The oldest dune group (Foxton Dunesand, see Cowie, 1963) laps on the post-glacial cliff and probably began to accumulate soon after the sea retreated from it as progradation began. 14C dates for woody peats interfingering with the oldest Foxton dunes at Waikanae are 4730 and 5290 years B.P. (N.Z. 2973) suggesting that the Waikanae coast was already prograding before the sea retreated from the cliff behind Centennial Inn, a situation comparable with the present. The next formation, Taupo Dunesand, formed largely from pumice granules derived from drift pumice piled on the beaches after the Taupo Pumice eruption (c. 1800 B.P.), is a prominent marker from
The table of formations (Table 1) summarises the stratigraphy and dates.
In Part I of these studies the problem of microbial contamination of aviation fuel was discussed and the role of the ‘kerosene fungus’ Cladosporium resinae (Lindau) de Vries (perfect state Amorphotheca resinae Parbery) in contamination and corrosion evaluated. Part II deals with the natural habitat of this interesting fungus.
Cladosporium resinae is not only capable of growing in and utilising kerosene as a source of food, it can grow on wood impregnated with coal tar products such as creosote and utilise these products as food (Christensen, Kaufert, Schmitz and Allison, 1942). In the late 1930's several people (Dr. C. et al., 1942). Because of the known toxicity of these preservatives to micro-organisms in general these workers initially more or less disregarded this sporadic appearance of a fungus on treated wood and thought of it only as a laboratory phenomenon. This is understandable when one considers that coal tar products, especially creosote, have been used for over one hundred years to protect wood from attack by fungi, insects and other organisms. According to Christensen et al. (1942), each cubic foot of wood impregnated with these products contains 8 to 20 pounds, or 1 to 2½ gallons, of the preservative and the long service of timbers so treated is evidence of the value of the coal tar products as preservatives. However, in the fall of 1939 an investigation was initiated by
Hormodendrum resinae (= Cladosporium resinae — see Part III) which Lindau had described in 1907 from the resin of a conifer, Picea excelsa, and that it is a common inhabitant of wood treated with coal tar products including creosote. These workers also isolated the same fungus from resinous wood and from asphalt street pavements.
Some years later Marsden (1954) became interested in this fungus in connection with its possible deleterious effects on the preservative used to protect the wood against decay. He confirmed the findings of Christensen and his colleagues regarding occurrence on creosoted timbers and deep penetration into wood. He found fruiting structures produced within tracheid cavities as well as on the surface of the wood and indications that this mould may deleteriously affect the toxicity of creosote. He described five strains of the fungus (see Part III), and he coined the name ‘creosote fungus’ in 1954. Christensen and colleagues were unable to isolate C. resinae from soil except on one occasion (glasshouse soil) and neither they nor Marsden were able to isolate it from the air. The indications were, therefore, that C. resinae occurred naturally in resinous wood and on creosoted timbers, but not in soil.
With the implicating of C. resinae in contamination of jet aviation fuels and in corrosion of aircraft tanks in the 1960's interest in this fungus revived. Leonard and Klemme (1961) believed that this fungus could be soil-borne because most other Cladosporia could be isolated from the soil. But it was not until the late 1960's that this was confirmed by Parbery (1967). After devising a method which successfully isolated C. resinae from soil he investigated its occurrence in soils from Australia, Britain and Continental Europe. On the basis of his findings he concluded (1969b) that C. resinae is a natural component of the soil microflora and is widely distributed in nature. His conclusions are supported by the results of recent work in New Zealand (Sheridan, Steel and Knox, 1971).
None of the earlier workers were able to isolate C. resinae from the atmosphere yet Christensen et al. (1942) and Hendey (1964) were, according to Parbery (1969b), still of the opinion that it is airborne. Parbery (1969b) referred to three independent trappings of this fungus from the atmosphere, thus supporting this view. Within the last few years Harvey (1971—personal communication) has isolated C. resinae from the air in Wales in very small quantities, but consistently. More recently, using a method which selectively isolates this fungus from the air (Sheridan and Nelson, 1971b), it has been possible to trap it consistently from the atmosphere over Wellington, New Zealand (Sheridan, Sheridan, Hoverd and Nelson, 1971; Sheridan, 1971).
C. resinae in airfield fuel systems in Australia. (Figures from Hazzard, 1963.) Black dot + ve Hollow dot —ve
From the foregoing it is quite clear that this extraordinary and intriguing, if not treacherous, fungus is very elusive. It has slipped through the fingers of the earlier workers when they have attempted to isolate it from soil and air. Results obtained by the use of new methods have given a better understanding of its distribution and role in nature but we will have to be patient until results from the wider application of these methods become available. In this paper the body of knowledge available at the present time on the occurrence of C. resinae in nature is reviewed in an attempt to determine its natural habitat.
The first report in the open scientific literature of the occurrence of Cladosporium resinae in hydrocarbon fuels appears to be that of Hendey in 1964. He reviewed the occurrence of this fungus in kerosene-type fuel storage tanks and fuel tanks of aircraft. Hazzard (1963) identified C. resinae in aviation kerosene in Australia in 1961 but his results are contained in a technical report which has not been widely circulated. In the same year Prince (1961) reported finding a Cladosporium sp. in jet fuel in the United States of America and his photographs indicate that it was probably C. resinae. As stated in Part I of these studies, contamination of kerosene-type fuels with
C. resinae was widespread in the 1960's — until recently it was much easier to isolate this fungus from kerosene than from soil and Hendey's application of the name ‘kerosene fungus’ to this organism is indeed appropriate. The accompanying map (Fig. 1) shows the distribution of C. resinae in airfield fuel systems in Australia. The figures are taken from Hazzard (1963) but the map is incomplete because of the difficulty of locating some of the airfields. Figures are not yet available for New Zealand, hence the distribution of C. resinae cannot be plotted. (For figures for Australia and California see Part I.)
The fungus has been isolated from lighting and aviation kerosene in France (Nicot and Zakartchenko, 1966 — ‘Mazout, kérosène carburants pour l'aviation’) and we have isolated it from both of these in New Zealand. We were not able to confirm from the literature that C. resinae had been isolated from petrol (= gasoline), diesel oil or lubricating oils, but the first sample of petrol we examined yielded five colonies of C. resinae from 250 ml. of the sample. Fuel/water samples of aviation gasoline are at present under test.
C. resinae has been isolated from aviation fuel, from tank filters or aircraft fuel tanks in Australia (Hazzard, 1963), Brazil (Gutheil, 1966), Denmark, England, India, Nigeria, New Zealand, Syria (Anon., 1968), U.S.A. (Engel and Swatek, 1966) and Japan (Inoue, Iwamato and Minoura, 1965).
It has been reported that C. resinae can remain alive for rather a long time in kerosene-type fuels (Hendey, 1964). Nicot and Zakartchenko (1966) found the fungus in their samples to be alive after 30 months but the results of their tests were not consistent. Hazzard (1967) found that it could remain viable for several years in essentially water-free petroleum products. This aspect has not yet been studied in our laboratory.
Four forms of C. resinae are known to exist (see Part III) but only one, j. avellaneum, has apparently been recovered from kerosene-type fuels (Hendey, 1964; Inoue, Iwamoto and Minoura, 1965) (Table 1). C. resinae f. resinae has arisen in culture from f. avellaneum (Hendey, 1964) and been isolated occasionally from soil (Parbery, 1968). An albino has arisen in culture from f. avellaneum (Parbery, 1969a; Sheridan, Steel and Knox, 1971) and is morphologically similar to it (Figs. 2 and 5). De Vries (1955) gave the name f. albidum to an albino which was morphologically similar to f. resinae. For the present purpose we shall refer to both of these as f. albidum (see Part III).
The only form isolated by us directly from kerosene fuels (and soil) is C. resinae f. avellaneum. Both f. albidum and f. resinae have been found in New Zealand as saltants in culture. All three forms have been tested by us for ability to grow in kerosene; all were able to do so (see Fig. 3). Parbery's (1968) isolates of f. resinae from
C. resinae f. avellaneum on V-8 juice agar containing 0.1% creosote.
Cladosporium resinae was repeatedly isolated from wood impregnated with creosote and coal tar, e.g. telephone poles and railway ties, by Christensen, Kaufert, Schmitz and Allison (1942). Samples were taken near the ground line from twenty-nine creosoted poles in Minnesota, Wisconsin, Pennsylvania and Delaware. C. resinae was isolated from twenty-seven of these. It was likewise isolated from six of eight creosoted ties sampled in Minnesota, and from all of eleven ties from Ohio, from all of four ties from Louisiana, three from the State of Washington, and from ties sampled in Missouri and Indiana. The fungus was also found on several creosoted southern pine fence posts in service for seventeen years in Iowa. These workers also successfully isolated this fungus from resinous woods.
Their work led the above workers to an investigation of the natural habitat of the fungus. Soil was tested around the University Farm, St. Paul, Minnesota, which included cultivated and uncultivated land, and soil near a creosoting plant but no C. resinae was found. Neither were they able to isolate it from the atmosphere. The reason for this failure to isolate the kerosene fungus from the soil and air was almost certainly due to the use of unsuitable techniques (see 3. The occurrence of C. resinae in soil). Marsden (1954) isolated
De Vries (1955) has studied various strains of Cladosporium resinae including two of Marsden's (1954). He concluded that Marsden's New York strain agreed with C. resinae f. avellaneum and his New York saltant strain with f. resinae (Fig. 5). The fungus isolated by Christensen et al. (1942) appears to be f. avellaneum (Table 2). De Vries tested the growth of five strains of the fungus on media containing 4% coal tar or 1% creosote. The strains tested were: 1, Enola; 2, ‘Christensen’ (received from Marsden); 3, C. resinae f. avellaneum; 4, C. resinae f. resinae; 5, C. resinae f. albidum. Contrary to De Vries's expectations, the last three strains did not grow on these media while the Enola and Christensen isolates made good growth. More recent work confirms that this fungus produces a wide variety of morphological forms and these vary in their ability to tolerate and grow in kerosene and creosote (Hendey, 1964; Parbery, 1969b). This aspect is dealt with more fully in Part III of these studies.
Cladosporium resinae was first isolated successfully from soil by Parbery (1967), working in Australia. He devised the ‘creosoted matchstick method’ in which two decapitated matchsticks, sterilised and soaked in sterile creosote, were added to a Petri dish containing moistened soil (See Fig. 4). The plates were incubated at 25° C. along with control plates containing non-creosoted matchsticks for up to three weeks. In Victoria, Parbery took soil samples from the base of, and some distance from, test creosoted poles and from virgin bushland, 50 metres from the testing site. C. resinae appeared in twelve out of fourteen soil samples tested. Since the fungus appeared in four samples of soil from the virgin bushland, this was considered by Parbery (1967) to be a strong indication that the fungus occurs naturally in soils. This observation led Parbery to further studies on the distribution of C. resinae in soil (Parbery, 1969b). He collected sixty isolates of C. resinae from soils in Australia. Britain, France and Sweden, proving the fungus to be widely distributed and indicating that it is a natural component of the soil microflora. The collection sites of soil yielding C. resinae were:
In England — fallow soil, under pines and poplar, by a creosoted pole, from soil under a larch and under Norway spruce.
In Wales — garden soil, the bases of creosoted poles, grass tussock and soil under firs and pines.
In France — soil under juniper, Norway spruce, oak, beech, Picea pungens, Cedrela sinensis and a soil bitumen mixture near base of pole.
In Sweden — there were two positive isolations from Stockholm, under Picea pungens and under oak and elm.
In Australia — isolates from Victoria were collected from burned hilltop forest, alluvial soil, beside a creosoted pole, skeletal hillside, roadside under eucalypt, petrol station, virgin bush, mountain soil of virgin bush, sandy arid soil and under Pinus radiata.
In New Zealand this fungus was first isolated from a garden soil in Wellington, near a creosoted fowl house, in November, 1969 (Sheridan and Knox, 1970). A modification of Parbery's ‘creosoted matchstick method’ was used. The matchsticks were steeped in creosote for 48 hours, transferred to a clean beaker covered with aluminium foil, and then autoclaved for 30 minutes at 103 kNm-2. This modification ensured that very little liquid creosote was transferred to the soil. This is an advantage because none of our isolates would tolerate as much as 3% creosote and the presence of excess creosote could suppress growth. C. resinae has subsequently been found to be widespread in soil in this country (Figs. 6 and 7), occurring most frequently in and near built-up areas but also occurring occasionally in soils remote from human habitation.
resinae. Left: f. avellaneum. Right: f. resinae.
We have isolated C. resinae in New Zealand from cultivated garden soils, from soils at the base of creosoted poles and fence posts and soils at the edge of a tar spillage on a roadside, from the top of sawdust mounds on old saw mill sites (Ketetahi Mills, near Taurewa, and near spiral railway, Raurimu), from soil under grass, gorse, conifers, scrub and native bush and from sandy soil in the Rangipo Desert. It has been found from sea level up to 5,000 feet. The collection sites are shown in Table 3.
Parbery (1969b) reports that there does not appear to be any correlation between isolation of C. resinae and any soil or vegetation type but there does appear to be some positive correlation between frequency of isolation and the annual average rainfall. This correlation could, in his opinion, be related to soil mosture or to organic matter content. This aspect warrants further study. In both Parbery's and our own studies not all soil samples taken beside creosoted poles yielded C. resinae, and several isolates were obtained from what is regarded as virgin soil.
C. resinae does not compete actively with other micro-organisms in a soil plate in the absence of creosote but this does not exclude it from being a soil inhabitor. Jones and Edington (1968) have demonstrated that there is an ecological niche available to hydrocarbon-utilising micro-organisms in soil. According to Parbery (1968) C. resinae can possibly utilise oils, waxes, steroids, terpenes, hydrocarbon and other biochemically related compounds in the soil for which it is able to compete against other micro-organisms. More recently Jones (1970) has reported studies on the origin and distribution
C. resinae in soils in New Zealand. Red spot +ve Black spot —ve
C. resinae in soils in Wellington area. Note position of Hirst spore trap. Red spot +ve, black spot —ve.
C. resinae in soils and the nature of hydrocarbons presents. While the distribution of this fungus in New Zealand suggests that the spread of C. resinae is associated with the construction of tar-sealed roads and the use of creosoted timbers in fencing and as power poles, the fungus has been found from time to time in areas remote from roads and from creosoted timbers. In some places at least, C. resinae must have been a natural component of the soil microflora for a very long time. In the near future it is hoped to examine soils from the old kauri forests of New Zealand for the presence of C. resinae. If it is found there this would be evidence to support the suggestion that C. resinae was present in this country before the advent of the jet age; perhaps before the arrival of the white man. This fungus grows readily on kauri gum so it is possible that it will be found in the old kauri forest areas.
Parbery (1969b) reports that of seventeen isolates tested, seven were able to grow in medium with kerosene as the only carbon source. All of the New Zealand soil isolates tested by us were able to grow in medium with aviation turbine kerosene as sole source of carbon but not all isolates from kerosene fuel could do so (Sheridan and Nelson 1971a).
Data on the occurrence of C. resinae in soil and the forms of the fungus found is presented in Table 4.
Christensen et al. (1942) exposed in their laboratory and out-of-doors, creosoted wood blocks and agar containing creosote but failed to recover C. resinae from the air. Nevertheless, they concluded that this did not exclude the possibility of spores being airborne or being present at certain times and places. Neither Marsden (1954) nor Hendey (1964) reported successful isolation from the air. According to Parbery (1969b), Christensen et al. (1942) and Hendey (1964) were of the opinion that spores of the fungus are airborne even though they failed to demonstrate this. Parbery (1969b) reports knowledge of three independent trappings of this mould from the atmosphere which support this view. These are: Anon. (1961) refers to a culture of C. resinae collected from an air filter, Chabert (1968) reports isolating it several times from the atmosphere over Rabat, Morocco, and Dr. H. J. Stewart, University of Melbourne, has given Dr. D. G. Parbery a culture of C. resinae collected from air over Johannesburgh, South Africa. Parbery (1969b) concludes that it is probable that the general failure to trap this mould from the air demonstrates the inadequacy of technique rather than the absence of spores of this mould in the air. Other experiments carried out
C. resinae did not remain viable when they were exposed to light (Hirst, 1971—personal communication).
Harvey (1967; 1971 — personal communication) has isolated C. resinae from the air in Wales in very small quantities, but consistently over a period of years. Parbery (1969b) was apparently not aware of Harvey's work.
We became interested in the possibility that C. resinae was a significant component of the airspora in New Zealand because of its widespread occurrence in soils and profuse sporulation, but all our initial attempts to trap it failed. After some ten months work we eventually perfected a technique which consistently isolated the fungus from the air of our laboratory. The application of the same technique out-of-doors has resulted in numerous trappings. A Hirst spore trap (see Fig. 8) is in use which is capable of continuous operation for twenty-four hours unattended.
Because C. resinae grows readily on a wide variety of agar media containing creosote (Sheridan, Steel and Knox, 1971) and few other fungi (or bacteria) will tolerate the creosote, we experimented with the use of a selective medium containing creosote for the isolation of this fungus. The method described here is a simple one and has proved suitable for isolating C. resinae from the air in New Zealand
(Sheridan and Nelson, 1971b). The selective medium is prep