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THE
PROC B DUNGS
THE
RINNE SOCErRY
OF
New SoutH WALES
FOR THE YEAR
1935
VOL. LX
WITH NINETEEN PLATES and 417 Text-figures.
SYDNEY: PRINTED AND PUBLISHED FOR THE SOCIETY BY
AUSTRALASIAN MEDICAL PUBLISHING CO., LTD., Seamer Street, Glebe, Sydney,
and SOLD BY THE SOCIETY. 1935.
CONTENTS OF PROCEEDINGS, 1935.
PARTS I-II (Nos. 257-258). (Issued 15th May, 1935.)
Pages. Presidential Address, delivered at the Sixtieth Annual General Meeting, 2th Marchs 1935: by, Professor IW. Ja Dakin, SDISe) 1.5) 5) Poe a eel xexexail
Elections bck geRbae Eat. Nato sasace | cate Wee tere SENS 2 aecri AROS MO y ANTS kt ee Xxxii Balance-sheets for the year ending 28th February, 1935 per ale ?.©0-0.01) J 9,0:0.07
Revision of Australian Lepidoptera. Oecophoridae. iii. By A. Jefferis TUNE VAD RSH SS Mea e au ee lesa MNS. | Breed green es on eaten 1-15
The Petrology of the Hartley District. iii. The Contact Metamorphism of the Upper Devonian (Lambian) Series. By Germaine A. Joplin, BISCyn .Gblateni sand) three exch) sain nyse eect 16-50
The Diptera of the Territory of New Guinea. ii. Family Tipulidae. By Charles P. Alexander. (Communicated by Frank H. Taylor.) GRwentyis Dext-feureseyie ae" bu ave fia" Gey Uae A eee ee 51-70
Australian Rust Studies. v. On the.Occurrence of a new form of Wheat Stem Rust in New South Wales. By W. L. Waterhouse PA ec 71-73
The Diptera of the Territory of New Guinea. iii. Families Muscidae and Tachinidae. By John R. Malloch. (Communicated by Frank H. Raylor)y COne: Text-femreny cick Ge? Seeks, ee ee 74-78
On some Australian and South African Species of Acarina of the genus Stereotydeus. (Penthalodidae.) By H. Womersley, F.R.E.S., A.L.S. (Three Text-figures. ) Fax eleid Gaels ia ered, ile) een a ee 79-82
Notes on the Mosses of New South Wales. ii. Additional Records. By Alan Burges, M.Sce., late Linnean Macleay Fellow of the Society in BOtamy dee co tees etek th odes rere RE es ee SUGs a GA SN 83-93
Studies in the genus Uromycladium. ii. Notes on the Dikaryon Stage of Uromycladium Tepperianum. By Alan Burges, M.Sc., late Linnean Macleay Fellow of the Society in Botany. (Sixteen Text-figures.) .. 94-96
An Investigation of the Sooty Moulds of New South Wales. iii. The Life Histories and Systematic Positions of Aithaloderma and Capnodium, together with descriptions of New Species. By Lilian Fraser, M.Sc., Linnean Macleay Fellow of the Society in Botany. (Sixty-five Text- figures. ) ee me a on renee NE G5 oo 97-118
The Gasteromycetes of Australasia. xvii. Some new Species of Hymeno- gastraceae. By G. H. Cunningham, D:Se:., Ph.D., H.R.S.N.Z. ., 9... «5 1a9=120
CONTENTS. ili
PARTS III-IV (Nos. 259-260).
(Issued 16th September, 1935.) Pages. The Relationship between Erosion and Hydrographic Changes in the Upper Murray Catchment, N.S.W. By Frank A. Craft, B.Sc., Linnean
Macleay Fellow of the Society in Geography. (Plates ii-iii and nine Text-figures. ) EME Ds ee, ee, Wi Oe ely Cee Ss oe PE RP eee MOA 4
Contributions to the Microbiology of Australian Soils. iii. The Rossi- Cholodny Method as a Quantitative Index of the Growth of Fungi in the Soil, with some Preliminary Observations on the Influence of Organic Matter on the Soil Microflora. By H. L. Jensen, Macleay
Bacteriologist to the Society. (Two Text-figures.) .. .. .. .. 145-154 Notes on Australian Orchids. A Review of the Species Dendrobium teretifolium R.Br. By the Rev. H. M. R. Rupp, B.A. (Plate iv.) .. 155-158
An Investigation of the Sooty Moulds of New South Wales. iv. The Species of the Hucapnodieae. By Lilian Fraser, M.Se., Linnean Macleay Fellow of the Society in Botany. (Ninety-one Text-figures.) 159-178
Australian Coleoptera. Notes and New Species. No. ix. By H. J. Carter,
B.A., F.R.E.S. (Seven Text-figures. ) Ee ee : . 179-193 The Marine Algae of Lord Howe Island. By A. H. S. Lucas, M.A., B.Sc. (Plates v-ix; seven Text-figures.) eee tah doimibelea Bie Nuatiak pale. une itl Nba yy
The Relations between the Internal Fluid of Marine Invertebrates and the Water of the Environment, with Special Reference to Australian
Crustacea. By Enid Edmonds, M.Sc. (Five Text-figures.) j6 og CBO ZEe Miscellaneous Notes on Australian Diptera. iii. By G. H. Hardy .. .. 248-256
Additions to our Knowledge of the Flora of the Narrabeen Stage of the Hawkesbury Series in New South Wales. By N. A. Burges, M.Sc. (Giatemxeandselevent wext=feunTeS)) ms eet eee en teen renee io. 04:
Upper Permian Insects of New South Wales. iii. The Order Copeognatha. By R. J. Tillyard, M.A., Sc.D., D.Se., F.R.S. (Thirteen Text-figures. ) SO Ee ae EE een RT ORIN Mea hi wl day Ay ara mir MA NaOCl
An Investigation of the Sooty Moulds of New South Wales. v. The Species of the Chaetothyrieae. By Lilian Fraser, M.Sc., Linnean Macleay Fellow of the Society in Botany. (Thirty-nine Text-figures.) 280-290
Notes on Australasian Anisopodidae (Diptera). By Mary E. Fuller, B.Sc. (GChirty-eicits Text-flZuTrest)r ess wich ates Tedke eS Ake cae tate se 291302
PARTS V-VI (Nos. 261-262).
(Issued 16th December, 1935.)
Observations on the Seasonal Changes in Temperature, Salinity, Phos-
phates, and Nitrate Nitrogen and Oxygen of the Ocean Waters on the Continental Shelf off New South Wales and the Relationship to Plankton Production. By W. J. Dakin, D.Sc., F.Z.S., and A. N.
Colefax, B.Sc. (Plate xi and eleven Text-figures.) oo od (ao tol OR Bie!
IV CONTENTS.
Pages. Revision of Australian Lepidoptera. Oecophoridae. iv. By A. Jefferis
Turner, M.D., F.R.E.S. é 315-339 The Leaf Anatomy and Vegetative Characters of the Indigenous Grasses
of New South Wales. i. Andropogoneae, Zoysieae, Tristegineae.
By Joyce W. Vickery, M.Sc. (Forty-three Text-figures.) 340-373 Upper Permian Insects of New South Wales. iv. The Order Odonata.
By R. J. Tillyard, M.A., Sc.D., D.Se., F.R.S. (Plate xii, figs. 1-3
and four Text-figures.) site| Haas, aS waSachin Fiscelg Al thee ESCO ce AG OTe Oe Upper Permian Insects of New South Wales. v. The Order Perlaria or
Stone-flies. By R. J. Tillyard, M.A., Se.D., D.Sc., F.R.S. (Plate xii,
figs. 4-5 and six Text-figures. ) 385-391 On the Climate and Vegetation of the Koonamore Vegetation* Reserve to
1931. By T. G. B. Osborn, J. G. Wood and T. B. Paltridge. (Plates
xili-xvii and ten Text-figures.) Eee eh i EE a CO ROO Cae Dele Studies in the Australian Acacias. v. The Problems of the Status and
Distribution of Acacia Baileyana F.v.M. By I. V. Newman, M.Sc.,
Ph.D., F.L.S., Linnean Macleay Fellow of the Society in Botany.
(Plate xviii and three Text-figures.) (With a Note on the Occur-
rence of Hybrid Acacias, by E. Cheel.) S88 eats : . 428-446 Note on the Permian Sequence in the Werrie Basin. With Description of
New Species of Fossil Plants. By S. Warren Carey, M.Se. (Four
Text-figures. ) ‘ .. 447-456 A Preliminary Note on the Acacia Legume as a Lateral Organ. By I. V.
Newman, M.Sc., Ph.D., F.L.S., Linnean Macleay Fellow of the Society
in Botany. (Six Text-figures. ) SA ee ae 457-458 Some Fossil Seeds from the Upper Palaeozoic Rocks of the Werrie Basin,
N.S.W. By A. B. Walkom, D.Se. (Plate xix.) 459-463 List of New Families, Genera and Subgenera .. 465 List of Plates 466 Abstract ofwProceediniesies |i ie a Syn a reer Pap SeXOGVATIT OX] ay alnn Donationswandelixchances: «ea. as anna a ee xlvili-lvili List of Members lix—lxiii Index lxiv-lxxv
CORRIGENDA (1935).
Page 92, line 2, for australis read australe for Hampella read Hampeella Page 111, lines 5, 6, for uniseptum read uniseptatum
Page 228, line 6 from bottom, for Bryopsis comosa read Bryopsis plumosa
Page 228, line 3 from bottom, for crassinervius read crassinervia Page 228, line 2 from bottom, for Helminthocladia read Helminthora Page 406, line 35, for Hrodium cygnodium read Hrodium cygnorum
Page 406, line 36, and Page 423, line 8 from bottom, for Tetragonia eremea read
Tetragonia eremaea Page 419, line 6, for Ah. Georgeii read K. Georgei
ANNUAL GENERAL MEETING. WEDNESDAY, 27th Marcu, 1935.
The Sixtieth Annual General Meeting was held in the Society’s Rooms, Science House, Gloucester Street, Sydney, on Wednesday, 27th March, 1935.
Professor W. J. Dakin, D.Sc., President, in the Chair.
The minutes of the preceding Annual General Meeting (28th March, 1934) were read and confirmed.
PRESIDENTIAL ADDRESS.
Another decade has passed in the history of the Society, which now enters its sixty-first year. Since the celebration of the Society’s Jubilee ten years ago there have been many changes—the most outstanding in the way of achievement perhaps being the co-operation with the Royal Society of New South Wales and the Institution of Engineers, Australia, which resulted in the building of Science House, now regarded by a number of the scientific and professional institutions of Sydney as their permanent home. During the same decade there have been startling changes financially, but the Society, in spite of one or two misfortunes, appears to have safely negotiated the worst of the bad times— chiefly as a result of the conservation of most of its surplus income in the prosperous years before the depression. We may justly be proud of the amount and quality of the research work carried out by members, which is placed on permanent record in the Prockgepines, of which the last ten volumes will compare favourably with those of any previous decade in the Society’s history.
The concluding part of Volume lix of the Society’s ProcrrpINGS was issued in December. The complete volume (447 plus Ixiv pages, nineteen plates and 351 text-figures) contains thirty-six papers from twenty-seven authors, five papers being by Linnean Macleay Fellows and three by the Macleay Bacteriologist.
Exchanges from scientific societies and institutions totalled 1,795 receipts for the Session, aS compared with 2,084, 1,866 and 1,703 for the three preceding years. During the year the following institutions were added to our exchange list: Imperial Fisheries Institute, Tokyo, Japan; New York Botanical Garden, New York, U.S.A.; University of Minnesota, Minneapolis, U.S.A.; Asociacion Sudamericana de Botanica, Montevideo, South America.
Since the last Annual Meeting the names of nine Ordinary Members have been added to the roll, three have been lost by death, three have resigned, and the names of three have been removed on account of arrears of subscription.
TANNATT WILLIAM HpGEWoRTH Davin, who died at Sydney on 28th August, 1934, was born at St. Fagan’s Rectory, near Cardiff, Wales, on 28th January, 1858. He was educated at Magdalen College School, and New College, Oxford, where he was elected to the Senior Classical Scholarship in 1876. He graduated B.A. in 1880. Included in his studies was a course of Geology under Professor Prestwich, and he made his first acquaintance with glacial problems in South Wales, his A
ii PRESIDENTIAL ADDRESS.
first paper, “Evidences of Glacial Action in the Neighbourhood of Cardiff’, being published in 1881 by the Cardiff Naturalists’ Society. He continued his geological studies under Professor Judd at the Royal School of Mines, and came to Australia in 1882 as Geological Surveyor on the staff of the Geological Survey of New South Wales, under the late C. S. Wilkinson. During the next decade he carried out many important geological investigations in New South Wales, including studies of the fossiliferous Silurian beds at Yass, the tinfields of New England, and the Coal Measures of the Hunter River district. The Hunter River Coalfield occupied much of his time and thought from this period until his death. During his survey of the field he discovered the occurrence of the Maitland Coalfield, and it is almost impossible to estimate the commercial value of the results of this work. Apart from the economic aspect, there arose many interesting problems concerned with the peculiar fauna and flora of the Permo-Carboniferous rocks, and with the occurrence of glacial phenomena.
In 1891 he was appointed Professor of Geology in the University of Sydney, where he remained until his retirement in 1924. His enthusiasm and inspiring personality quickly widened the influence of the Geological School of the University, and under him there grew up a band of geologists who have taken a prominent part in the development of geology and mining in Australia. He soon became recognized as a leader amongst Australian scientists; he was President of the Geological Section of the Australasian Association for the Advancement of Science in Hobart in 1892, and Brisbane, 1895; President of our Society, 1893-4 and 1894-5; President of the Royal Society of New South Wales, 1896 and 1909; President of the Australasian Association for the Advancement of Science, 1904, at Dunedin, and 1913, at Melbourne; President of the Australian National Research Council, 1921-22. He was a member of the Council of our Society from 1891 until his death, and was also for many years a member of the Council of the Royal Society of New South Wales.
His presidential addresses form a valuable series of summaries of knowledge in the Australian Region of volcanic action, structural features, evidences of glaciation, Mesozoic History, and tectonics.
In 1897 he was chosen as leader of the second expedition to the Atoll of Funafuti, where, in addition to obtaining a complete core from a bore sunk to a depth of 1,118 feet, he carried out a survey of the atoll ‘and made investigations on the growth of corals. Soon after his return he studied the great thickness of Radiolarian rocks of Devonian age in New South Wales, showing that they were laid down in comparatively shallow water and not in abyssal depths. In 1906 he visited the glaciated districts of Southern India and attended the International Geological Congress in Mexico, where he presented an important paper summarizing the hypotheses put forward to explain past changes in climate. The year 1908 he spent in the Antarctic with the Shackleton Expedition, making the first ascent of Mount Erebus, and also made the first journey to the South Magnetic Polar area.
Much of his time for a few years after his return was occupied in arranging for the study of the geological material brought back by the Expedition, and also in securing funds for the publication of the scientific memoirs of the Expedition, this latter involving lecturing tours throughout the Commonwealth. Then came further Antarctic activities—organization of the Australasian Antarctic Expedition under Douglas Mawson, arrangements for Captain Scott’s last Expedition, and securing support for Shackleton’s Second Expedition—and the
PRESIDENTIAL ADDRESS. iii.
visit of the British Association to Australia in 1914, in all of which he took a prominent part. On the outbreak of war he took an important part in the organization of a battalion of miners; he himself joined as Major, and arrived in France in May, 1916. He became geological adviser to the Controller of Mining in the First, Second, and Third Armies, and later to the Inspector of Mines of the British Expeditionary Forces, and in this capacity rendered very valuable service, since geological advice was of the greatest importance in tunnelling and mining operations in the very porous strata below ground water level. He was promoted to the rank of Lieutenant-Colonel, received the D.S.O., and was twice mentioned in dispatches.
In 1924 he retired from the Chair of Geology to devote himself to the preparation of a work on the Geology of Australia. He supplemented his previously unrivalled knowledge of the geology of the continent by travelling extensively and visiting or revisiting many critical areas. In 1933 he published a new Geological Map of Australia, accompanied by a volume of explanatory notes, but unfortunately he had not completed the major work at the time of his death. He spent much time during his last few years in studying the traces of the remains of organisms in the Pre-Cambrian rocks of South Australia. Though he had not yet succeeded in convincing all of his colleagues that the remains were truly organic, he himself believed that this piece of work was one of the most important contributions, perhaps the greatest, he had made to science.
As a geologist and as a leader in science his fame was world-wide, and he received many honours in recognition of his outstanding achievements: He was made C.M.G. in 1910, D.S.O. in 1918, and K.B:E. in 1920; he was awarded the Bigsby Medal (1899) and the Wollaston Medal (1915) of the Geological Society of London, the Conrad Malte-Brun Prize of the Geographical Society of France (1915), the Mueller Medal of the Australasian Association for the Advancement of Science (1908), and the Clarke Memorial Medal of the Royal Society of New South Wales (1919). He had conferred on him the honorary degree of Doctor of Science by the Universities of Oxford, Wales, Manchester, Cambridge and Sydney, and of Doctor of Laws by the University of St. Andrews. The Royal Society of London elected him a Fellow in 1900. On his retirement in 1924 he was made Professor Emeritus by the Senate of the University.
He was truly “a fine scholar, a great scientist, a gifted teacher, a distinguished explorer, an ardent patriot, a warm-hearted philanthropist, a gracious friend, and a humble-minded Christian gentleman”.
WALTER H. Bone, who died at Killara on 15th July, 1934, had been a member of the Society since 1923. He was a great lover of the bush and a writer of animal and bush stories. As a naturalist he was a supporter of various scientific societies, but it was only on very rare occasions that he attended meetings of our Society.
Tuomas McDonnovueH died at his home at Coogee on 26th June, 1934, at the age of sixty-seven. He entered the Public Service in 1891 and was engaged on the Sydney Detail Survey until he was transferred to Ballina in 1901. Here he spent several years in survey work in connection with water supply, drainage schemes, and harbour and river works; in 1908 he was transferred back to Sydney, and for many years was occupied in investigations for sewerage and water supply schemes for a number of the larger country towns of New South Wales, as well as for drainage and sewerage works in Sydney. He was a licensed surveyor and an Associate of the Sydney Technical College. Though he did not
iv PRESIDENTIAL ADDRESS.
take an active part, he was for many years a very regular attendant at the meetings of this Society, of which he had been a member since 1907.
WILLIAM SUTHERLAND DuN, who died at Mosman on 7th October, 1934, was born at Cheltenham, England, on ist July, 1868. At the age of about twelve months he came to Australia on the ship ‘Sobraon’. He was educated at Newington College and the University of Sydney, and entered the Department of Mines in 1890. In the earlier years of his service in the Geological Survey he was assistant to the late Sir Edgeworth David in the survey of the Hunter River Coalfields. Later he became assistant to the late Robert Etheridge, Jr., under whom he obtained his training as a palaeontologist. In 1899 he became Palaeontologist and Librarian to the Geological Survey, a position he retained until his retire- ment from the Public Service: in 1932. He was also Lecturer in Palaeontology in the University of Sydney from 1902 till his death, and was Honorary Palaeon- tologist to the Australian Museum. He was President of this Society for the two years 1913 and 1914, and a Member of Council from 1901 to 1919. He was an Ordinary Member of the Society from 1894 to 1922, and a Corresponding Member, 1932-1934. He was President of the Royal Society of New South Wales in 1916, and for many years a member of the Council of that Society. He was elected an associate member of the Australian National Research Council in 1922. He contributed only three papers to our ProcrEpines, one of them in conjunction with W. N. Benson and W. R. Browne, and one with W. H. Rands- and T. W. EH. David. The greater part of his published work was palaeontological, and much of it appeared in the publications of the Geological Survey of New South Wales. He had an extraordinarily wide knowledge of the fossil faunas and floras of Australia and of geological literature in general, and it was seldom that he was unable to assist any one of his fellow workers seeking information. His death was a very severe loss to Australian palaeontology, and, with the present-day tendency to specialization, it will probably be a very long time before another acquires such a wide knowledge of our fossils.
We offer our hearty congratulations to Mr. E. C. Andrews on his election as an Honorary Fellow of the Royal Society of New Zealand; Dr. R. J. Tillyard on the award of the Mueller Medal by the Australian and New Zealand Association for the Advancement of Science; Professor W. J. Dakin on the award of the R. M. Johnston Memorial Medal by the Royal Society of Tasmania; and Mr. John Andrews on the award of a Rockefeller Scholarship to enable him to continue his studies at Cambridge.
The year’s work of the Society’s research staff may be summarized thus:
Mr. H. L. Jensen, Macleay Bacteriologist to the Society, completed his intro- ductory work on the numbers of microorganisms in soils and the preliminary tests of the usefulness of the Rossi-Cholodny method. The results of this work appeared in two papers in the Procrepines for 1934. He then carried out preliminary experiments on the influence of varying temperature and moisture on the composition of the soil micro-flora in decomposition experiments with organic matter in soil. These experiments showed that, as a general rule, irrespective of the character of either the soil or the organic material in it, bacteria tended to multiply most strongly at low temperatures, whereas actino- mycetes predominated under conditions of high temperature and low moisture. By means of an adaptation of the Rossi-Cholodny method he was able to obtain quantitative expressions for the density of vegetative fungal mycelium in the soil, a method for which had hitherto been wanting. These results have been incor-
PRESIDENTIAL ADDRESS. Vv
porated in a paper which is ready for publication. He then commenced the main experiments on the decomposition of organic matter by soil organisms, the general aim of which was to determine over a period of ten to forty days, at temperatures varying from 38-5 to 37-38° C., the production of carbon dioxide from soil either with or without extra addition of organic matter, and to correlate the rate of carbon dioxide formation (which serves as an index of the intensity of decom- position) with the changes taking place in the numbers of the different groups of microorganisms. The results of these experiments have been consistent, and they account naturally for the rapid disappearance of “humus” in soils in hot climates as well as for the synthesis of proteid material which has been shown to take place during the decomposition of organic matter at low temperatures. They would also seem in part to account for the vigorous nitrate formation that is known to take place in Australian wheat soils, but more work in this direction is needed before any final conclusion can be reached.
Mr. F. A. Craft, Linnean Macleay Fellow of the Society in Geography, has completed two papers, one of which, “Regimes and Cyclical Volume Changes of the Upper Murray and Snowy Rivers, N.S.W.”’, appeared in the ProcrErpines for 1934. The second deals with the relationship between stream flow and modern erosion in the upper Murray Catchment. He has found that the work of the stream since white settlement has been directed towards the continuation of terrace cutting in valley alluvials. A cycle is indicated involving pebble formation from weathered channels and hillsides, followed by a period of decreasing supply of material and its more complete reduction in the passage of gorges and ending in clear streams flowing on unweathered rock. He also completed work for a further paper dealing with stream geography in south-eastern Australia. This has been mainly devoted to a preparation of a series of maps showing the relative annual flows of most of the principal rivers, the distribution and importance of effective catchments, the annual regimes, the variability of flow from year to year, and the importance of exceptional maximum and minimum flows in the output of streams.
Miss Lilian Fraser, Linnean Macleay Fellow of the Society in Botany, completed a study of the life-histories of Aithaloderma ferruginea and A. viridis, the results being inciuded in a paper on the life-histories and the systematic positions of Aithaloderma and Capnodium. These two genera are shown to be closely related, and the affinities of the Capnodiaceae prove to lie with the Dothideales rather than with the Sphaeriales or Perisporiaceae. It is shown that there may be developed structures such as “ostiole”’, periphyses and stromatic wall, which are very similar to structures in unrelated genera, but are of different origin. The importance of this in the systematic determination of mature specimens without examination of the life-history is discussed. Systematic studies have been made of (i) the species of Hucapnodieae collected in New South Wales, of which five species and varieties are described as new and other species are described, several of them being recorded from Australia for the first time; (ii) the species of the Chaetothyrieae collected in New South Wales, of which nine are described as new and one recorded for the first time in Australia; (iii) species of Meliolineae and Trichopeltaceae from various parts of New South Wales. Work in progress includes a survey of the host range, distribution, nomenclature and biology of Asterella Hakeae, and a microchemical study of the cell wall of Dematium pullulans. During the coming year Miss Fraser proposes to make a complete study of the cell membrane of the Capnodiaceae and related
vi PRESIDENTIAL ADDRESS.
fungi to determine, if possible, the reason for their powers of resistance to variations in temperature, humidity and light intensity. She also hopes to complete the study of the reactions of members of the Capnodiaceae to substances in the honey-dew of insects, with a view to discovering the reason for their restricted habitat. She also proposes to study the distribution of the epiphyllous flora of the rain forest areas and to collaborate with Dr. McLuckie in the description of new parasitic fungi.
Dr. I. V. Newman, Linnean Macleay Fellow of the Society in Botany, prepared for publication the later portion of the life-history of Acacia Baileyana _(Coota- mundra Wattle). He conducted extensive field work in a search for the natural habitat of this species, but was able to find it only in one small area near Cootamundra. Following on the phenomena of fertilization recorded in Acacia Baileyana, he collected. material for a further study of those phenomena in A. Baileyana and A. discolor. He has studied A. longifolia and A. suaveolens in view of the theory of carpel polymorphism, enunciated by Miss EH. R. Saunders of Cambridge, and has shown that these two species do not conform to that theory, as claimed by Miss Saunders. This study deals with the whole course of the ontogeny of the legume up to the time of fertilization, and will have some bearing on theories propounded by Professor J. McLean Thompson and Dr. H. Hamshaw Thomas. Dr. Newman has also begun a genetical study of the flower-colour forms of A. discolor, and has carried out a considerable amount of field and herbarium work for the future revision of the taxonomy of the genus Acacia. During the coming year he proposes to continue his investigation of the Australian Acacias with a view to working out a classification that will corres- pond with the phylogenetic relationships and will clarify the many difficulties existing in the taxonomy of the genus.
Mr. N. Alan Burges, Linnean Macleay Fellow of the Society in Botany, resigned his Fellowship as from 31st July, 1934, having been awarded the James King of Irrawang Travelling Scholarship by the University of Sydney, under which he proceeded to Cambridge. During the time he held the Fellowship he continued his study of Uromycladium, particularly that of U. Tepperianum on Acacia stricta. Material was collected in the field and some infection experiments were tried, but were unsuccessful. He continued the cytological examination of the teleuto- spore stage, paying particular attention to the origin of the binucleate stage. The results of his work have been embodied in papers which will be submitted to the Society during the coming year.
Three applications for Linnean Macleay Fellowships were received in response to the Council’s invitation of 26th September, 1934. I have pleasure in reminding you that the Council reappointed Miss Lilian Fraser and Dr. I. V. Newman to Fellowships in Botany, and also appointed Mr. R. N. Robertson, B.Sc., to a Fellowship in Botany for one year from ist March, 1935. We wish them a successful year’s research. The small number of applications was due, partly, to the fact that at least two prospective applicants received appointments of a more permanent nature very shortly before the time for application for Fellowships.
Mr. Rutherford Ness Robertson graduated in Science at the University of Sydney with first class honours in Botany in March, 1934. He was then awarded a Science Research Scholarship in the University. He has been investigating the physiology of the movement of stomata in certain Australian plants, and has come to the conclusion that this movement is intimately bound up with the
PRESIDENTIAL ADDRESS. vii
general metabolism of the leaf and is not specially connected with water loss. He has designed a special apparatus for extracting the gas from the intercellular space system of leaves and for analysing this gas for its relative percentage of carbon dioxide and oxygen at different times during the day. By this he hopes to discover any correlation which may exist between stomatal movement and photosynthesis and respiration. In addition to this physiological work, he has taken part in two ecological surveys. For his year’s work as a Fellow he proposes to continue the investigation of the physiological processes involved in the leaf, and particularly their bearing on stomatal movement. He also proposes, as opportunity offers, to continue his participation in the ecological survey of the Myall Lakes area and to elucidate some of the problems that arise there.
THe Aquatic ANIMAL AND ITS ENVIRONMENT. From the Point of View of Salinity and Osmotic Pressure of the Internal Media.
During the last ten years Zoological Science has made an altogether new valuation of the study known since 1869 as Animal Heology. The intricacy of the relationships between an animal and its environment (whether inanimate or animate) has been better appreciated. As a result there has arisen a realization of the need for a thorough investigation of all the mechanisms which subserve this relationship. The new Ecology will certainly provide future zoologists with enough experimental work to satisfy the most ardent critic of purely descriptive work. It will entail not only laboratory experimentation, but the closest observa- tion of the whole organism in its natural habitat and experimentation in this environment. And it will, no doubt, provide results of as much value in economic science as in pure science.
Tonight I propose to put together a story of the investigation of one aspect of this matter—the relationship existing between an aquatic animal and its environment on account of the fact that some, at least, of its bounding membranes must be permeable wholly or partially to the external medium (whether it be sea or fresh water).
Twenty-six years have actually elapsed since the publication of my first paper on this subject. At that time very few people indeed, in Great Britain, were interested in the matter. Earlier work had been carried out by several European scientists, a Canadian physiologist and a United States zoologist. Practically no further interest was shown in the British Empire until recent years. Perhaps the modern trend to make zoology a more experimental science is responsible for the new enthusiasm in this line of research which, one might add, has been almost newly discovered by zoologists. For this reason, I felt that it would be useful if, without entering into too much detail (surely quite an unnecessary feature of a Presidential Address), I set forth the position reached and something of the interesting tale of progress.
As far back as 1859, the great French physiologist, Claude Bernard, with that foresight so characteristic of him, realized the advantage of the term ‘Internal Medium” for those constituent fluids of the body (blood, coelomic fluid, etc.) as contrasted with what is outside. By the term Haternal Medium I shall under- stand the fresh water, sea water, hot spring water or whatever may be the aqueous fluid in which aquatic animals live. The term ‘Internal Medium” or “Internal Media” is a particularly useful one, for it serves to include other fluids of the body as well as blood, and in many invertebrates these body fluids play a very important part in the constitution of the body.
viii PRESIDENTIAL ADDRESS.
Now it might appear to be obvious that the fluids in the vacuolated protoplasm of a single-celled organism like an Amoeba, a Paramecium, a marine Radiolarian, or a multi-cellular creature like a Jelly Fish, should be much more related in constitution to the external medium than those of a crab with its hard shell of impervious chitin, or a fish with its scaly exoskeleton. The facts show, how- ever, that the real conditions are by no means so obvious and simple as one might conclude at first sight. The blood of a teleost fish is usually altogether different in its salinity from that of a lobster living side by side in the same sea-water. The thin protoplasmic membrane of an almost microscopic single cell may separate fluids astonishingly different in composition.
The most important difference between waters in which aquatic animals live lies in their salt content. We shall, therefore, be chiefly concerned with the effect of this saline composition of the external medium upon the creatures living within it. Fortunately the saline composition of ocean water is remarkably uniform. The degree of salinity varies in different piaces, and it may be very low in river estuaries, but even here one usually finds the different constituents in the same proportions. The only change has been a dilution. It will be desirable to give the constituents of a typical ocean water at the outset.
Chlorine .. oes ce 35 ae Hs ao 55:29 Bromine 0:19 SO, 7:69 COsee aN ae ede aS An oe 0-21 Sodium ae ae 8 bie — Be a 30°59 Potassium .. 1-11 Calcium a3 a a a ae he 1-20 Magnesium ae kf ae ae a a 3:72
The total weight of the salts in grams per 1,000 grams of sea-water is known as the salinity, and the average salinity of typical ocean sea-water may be regarded as 35%c.
Chemical analyses of the internal fluids of the animal body date back to the *fifties, but some of the early work was very inexact. Thus, in 1852, Thomas Williams stated that the bulk of the fluid of the visceral cavity of Tubularia consisted of sea-water, “for when the specimen dries and the fluid evaporates, cubie crystals of chloride of sodium are seen amidst the albuminous molecules”. The same author also considered that the bulk of the fluid in the peritoneal space of the Gephyrea was salt water.
The first more detailed and accurate chemical analyses of the fluids of aquatic animals came in the years between 1870 and 1888. The salinity of several of these was set out by Boussingault in 1872. Leon Frédéricq examined Octopus blood in 1878, and L. Cuenot investigated the starfish in 1888 and claimed that the body fluids were practically sea-water with all its salts.
It is evident that, about the period between 1880 and 1885, the relations between the internal media and the external media were beginning to be under- stood, and this was undoubtedly due mainly to the work of Leon Frédéricq of Liége. In a paper in 1882, bearing the title ‘Influence du milieu extérieur sur la composition saline du sang chez quelques animaux aquatiques”’, he states definitely that the blood of crabs, lobsters and octopus of the North Sea is as salt as the sea-water, whilst that of the crayfish of the rivers contains very little salt. And he adds to this the comment: “It seems, therefore, to be established that by virtue of the simple laws of diffusion an equilibrium in salts is produced by a simple exchange.” Then, however, comes the more interesting further state-
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ment to the effect that in fish the conditions are not like this and that, despite the fact that oxygen and CO, easily pass through the gills of these animals, the fish of the sea present a salinity entirely different from that of the water they live in.
Three years later, in 1885, Claude Bernard was fully acquainted with the consequences of these discoveries, and in his book “Introduction a l’étude de la médecine expérimentale” (p. 110) he proclaims: “Chez tous les étres vivants le milieu intérieur, qui est un produit de lorganisme, conserve des rapports nécessaires d’échange et d’équilibre avec le milieu cosmique extérieur, mais a mesure que lVorganisme devient plus parfait, le milieu organique se spécifie et s’isole en quelque sorte de plus en plus du milieu ambiant.”
It was a remarkable generalization, seeing that the real facts were only divulged here and there and in no case in a really complete state. Claude Bernard had realized that in the vertebrate phylum there had been evolved a remarkably constant internal chemical environment culminating in the regulated temperature of the birds and mammals.
Frédéricq, in commenting on Bernard’s generalization in the year of its publication, gives further details of his own work on the salinity of the blood of aquatic animals under different conditions, and now begins to try the experiment of putting a marine crustacean into a mixture of sea and fresh water. It is clear, however, that very little was known of the exact saline composition of the blood. It was the general concentration of salts that had aroused interest.
But already, from another side, facts were being obtained which were to have a very fundamental bearing on the question. For many years it had been customary for physiologists to use a solution of common salt (NaCl) when making experiments in which blood had to be diluted without the corpuscles changing in volume, or for the examination of fresh animal tissues under the microscope. The solution used was about 0-75%, approximately isotonic with the blood. In 1882 Sydney Ringer, in a famous paper, showed that if he desired to keep a frog’s heart beating the ordinary saline solution was disastrous. He then tried adding other substances to the normal saline in order to obviate the abnormal effects, and discovered that white of egg would do it. He traced this to the effect of potassium chloride, and eventually by a series of thoughtful experi- ments a solution was obtained which would maintain the heart beat satisfac- torily. The solution has since been known as Ringer’s Solution. It consists of NaCl 0:65%, KCl 0:03%, CaCl. 0:02% + a trace of sodium bicarbonate, and it is a curious fact that the relative proportions of sodium, calcium and potassium in this mixture are very close to the proportions of the same salts in sea-water.
In the year 1889 Bunge made the suggestion that the large amount of sodium chloride in human tissues might be the relic of some aquatic ancestor. A few years later, Quinton (1897), better versed now in the composition of the internal media of different animals, made the definite assertion that the internal medium was practically a marine medium and that even a highly developed creature, such as a bird or a mammal, should be able to withstand a considerabie introduction of sea-water. Finally, he extended his thesis and stated definitely that the facts pointed to a theory that life originated in water and that there could be no doubt that such water was marine.
There were now two theses in the field arising from a study of the salinity of the internal fluids of aquatic and other animals, and either of these alone was of sufficient interest to make the matter worthy of general attention.
B
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Singularly enough they were to remain practically unknown, except to the few specialists who had taken the subject up.
The idea that evolution had resulted in a progress from a condition in which the internal fluids of aquatic animals were entirely at the mercy of the external medium, to one in which the animal controlled its internal media and kept them independent of their surroundings, led naturally to a more detailed and accurate series of researches.
The questions to be answered were:
(1. In what animal groups did the independence of the internal media first become obvious?
2. What was the mechanism involved in maintaining this independence?
In regard to this second question, which was to prove by far the more difficult of solution, several possibilities could be envisaged. The skin and outer bounding membranes of the body could be impermeable to water and salts (as in the whales which live in sea-water but come to the surface to breathe); the skin could be permeable or semi-permeable and the regulation of the composition of the internal blood, ete., could be maintained by the kidneys or other excretory organs; the outer membrane itself could play a part in regulation.
The collection of more observations was clearly the first need, and several authors now commenced to make observations by other methods and more careful chemical analyses. :
Realizing that the skin and body wall of aquatic animals could act as a semi- permeable membrane with resultant osmotic conditions, Bottazzi of Naples commenced, in 1897, a series of investigations on the so-called osmotic pressure of the blood of fishes and the internal media of other aquatic animals. The principal method used was the determination of the freezing point of the fluids in question by use of the Beckmann Freezing Apparatus. This method was introduced about 1892 by Dreser for the investigation of human body fluids for medical purposes. :
Its application to the new line of research was particularly appropriate, for, not only is it a very convenient method, but it had the advantage of throwing light from a different angle on the relationships of the internal and external media.
The phrase “osmotic pressure” implies, of course, the presence of two selutions separated by a semi-permeable membrane. To speak, then, of the osmotic pressure of a fluid apart from these conditions may seem strange. Actually no difficulty arises in practice. When salts are dissolved in water and the solution is separated from pure water by a membrane impermeable to the salts but. permeable to the water, water passes through the membrane from the solution to the water and with a pressure which is dependent upon the concentration of molecules, ions or colloidal particles in the solution. When we speak of the osmotic pressure of a solution we refer, therefore, to the effect it would produce if it were separated by a semi-permeable membrane from the pure solvent. Now, since the freezing point of a solution is also lowered proportionately by the concentration of molecules, ions, etc., within it, we can use the freezing point as a direct measurement of osmotie pressure. Generally it suffices to give the lowering of the freezing point, thus salt water of 37-83%, freezes at —2:29° C. We express this as A 2-29°, meaning that the saline constituents are responsible for a depression of the freezing point of 2:29° C.
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We may now give some of the results of Bottazzi’s investigations in the
‘nineties. Coelenterata.
Alcyonium palmatum Vascular cavity fluid .. A 2:195 Echinoderms.
Asteropecten aurantiacus Water vascular system A 2:31 Asterias glacialis Visceral cavity fluid A 2-29 Gephurea.
Sipunculus nudus Fluid of visceral cavity A 2:31
Crustacea.
Maja squinado Blood A 2:36 Homarus vulgaris Blood A 2-292 Gasteropoda.
Aplysia limacina Body cavity fluid A 2°31
Cephalopoda. Octopus macropus Blood A 2-24 Elasmobranch fishes. Torpedo marmorata Blood A 2-26 Trygon violacea Blood A 2-44 > Marine Teleost fishes. Charanz puntazzo Blood A 1:04 Cerna (Serranus) gijas Blood A 1-034
The freezing point of the Naples sea-water from the Aquarium is given as having an average A of 2:29. The freezing point for human blood is —0:56 to — 0-59, and it is interesting to note that Rodier (1899) obtained A 0-602 for the turtle Chelonia caouana ana A 0:74 for the blood serum of a dolphin, Delphinus phocaena.
These figures bore out the analyses of Frédéricq (1884, 1885, 1891). The resemblance between the body fluids of the invertebrata and the sea-water in which they were living was clearly brought out. But a surprising feature was the low freezing point for elasmobranch blood, which indicated an apparently high salinity. It looked at first sight as if the sharks and rays resembled the inver- tebrates in the condition of their body fluids. Here was a really astonishing fact which was rendered more interesting still when Rodier, working at Arcachon, where the water was slightly less saline, found that the blood of six species of sharks and rays had a freezing point slightly different from those of Naples, but agreeing in that it was again nearly, if not the same as, that of the sea-water.
But Frédéricq (1891) had already shown that the marine elasmobranchs, like the teleosts, were relatively poor in salt. It seemed as if there were some curious discrepancy between freezing point determinations and analyses. The solution of the mystery was grasped by Quinton and Rodier (1899), both of whom realized that an extraordinary proportion of urea, already noted as a character of shark blood, was responsible for the unusual lowering of the freezing point.* Unfortunately, up to this time, the methods of analysis had been only approximate and the full situation was still not realized.
*'The fact that sharks and rays were apparently different from all other animals in containing an enormous quantity of urea in the blood was discovered as early as 1858 by Staedeler. In 1888 Krukenberg confirmed this for a number of species.
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In the meantime experiments were being conducted upon the effects of fresh- water or diluted sea-water on marine fish and other marine creatures, and vice versa, of sea-water on fresh-water forms. It was easily seen that an alteration in the sea-water by diluting with fresh-water produced a fall in the salinity of the internal media of many invertebrates, with a corresponding reduction in osmotic pressure.
A few of these experiments, combined with the results of the determinations made on animals from natural environments of different type, caused Bottazzi (1908) to put forward the argument that the body fluids of marine invertebrates, and also those of the elasmobranchs, had the same osmotic pressure (except for slight and unimportant differences) as the surrounding sea-water. The actual electrolytes present in the fluids of the different species varied, and might differ even in the same groups of animals. On the other hand, the internal media of marine teleosts differed entirely from the surrounding medium in osmotic pressure, and in this respect the teleosts resembled the higher land vertebrates.
But the invertebrates of fresh and brackish waters had to be regarded as entirely different from their marine relatives. The salt content of their blood was found to be higher than that of the surrounding water—in the case of the fresh-water crayfish much higher. We shall see that the conditions are altogether more complex here than was realized, and I shall return to the invertebrata after a consideration of the conditions in teleost fishes.
The Osmotic Conditions Prevailing in Teleosts.
There was considerable doubt at first as to the constancy and complete independence of the blood and internal media of the teleost fishes. Quinton, by putting fishes from sea-water into fresh-water, had indeed altered the concentration of the blood salts, but the fish were anything but normal under such conditions.
Garrey (1904) believed that the body membranes of teleost fishes were definitely impermeable. He examined a species of eel in both fresh and salt water and also the small fish so commonly used for experiments in the United States, Fundulus heteroclitus. He stated: “From these experiments we may conclude that in all probability the blood of Fundulus does not suffer much, if any, change in concentration when the fish is transferred from salt-water into fresh-water or vice versa, provided the membranes are uninjured. If these experiments admit of general application to migratory teleosts they would indicate that these animals also are in some way protected from changes in the osmotic pressure of the blood and tissues and that the principal protective factor probably lies in a lack of permeability of their membranes.”
Griffiths (1892), who had made chemical analyses of the body fluids and tissues of many species of invertebrates, stated also that the blood of a marine haddock did not contain more soluble salts than that from fresh-water fishes.
Naturally interest was directed to such fish as could pass normally from sea-water to fresh-water and safely withstand extensive changes in the constitution of the external medium. In this regard Greene (1904) investigated the Chinook Salmon for the United States Bureau of Fisheries, remarking that it might be considered an ideal subject for the study of the osmotic balance which existed between the outside medium and the living tissues.
Greene examined eighteen salmon from the sea and found that the mean freezing point of the blood was —0-762° C. He then examined salmon from tidal waters which were practically fresh and, finally, others from the spawning
PRESIDENTIAL ADDRESS. Xili
beds in fresh waters. The figure for the tidal waters was -—0:737 and for the spawning beds —0:613. Notwithstanding his determinations, Greene was averse to putting these results down as a proof of the effect of the surrounding water. He affirms that “the absence of food and the important metabolism occurring during the eight to twelve weeks’ sojourn in fresh-water are to be considered in this connection and possibly are sufficient to account for the change’’.
I am giving rather full attention to the matter of the fresh-water teleostei because it seemed that this was the lowest group of vertebrate animals which had achieved a full measure of control over the composition of their blood. It was essential to determine whether complete independence had been attained.
Some evidence was already accumulating against the view set out above, Dekhuyzen (1905) in particular having examined different teleost species from sea-water of a wide range of salinity. I was rather against any further attempt at aquarium experiments since the more or less rapid changes of water seemed extremely likely to upset the normal condition of the fish. And if the teleosts performed work in sustaining a blood salinity and osmotic pressure very different from the external media, it would be most likely that any unhealthy conditions would completely overshadow normal changes, especially if such were small.
Cruise of “Poseidon”, Feb.-March, 1908.—The stretches in black
connected by the dotted line indicate the places where the trawl
was used, and the numbers the series of determinations referred to in the text.
The method which I adopted in order to investigate the matter more fully was to obtain permission to travel on one of the scientific voyages of the German research ship “Poseidon” from Kiel in the Baltic to Helgoland in the North Sea. In this way it was thought possible (provided that trawling was carried out at
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regular intervals) to obtain fish of the same species—Plaice (Pleuronectes platessa) and the Cod (Gadus morrhua) in particular, as well as others—from water of a range of salinities, to obtain them in the living state, and to do all this within a few weeks, so that conditions might remain as uniform as possible during the entire experiment.
Since the account of this expedition was printed in the first numbers of a journal which is now somewhat rare, I have taken this opportunity to redraw the map of the voyage, which shows where the otter trawl was used and the actual determinations were made.
‘ The results which concern us here may be tabulated as follows:
Series A of Sea A of Date. No. Position. Water, Salinity. Species of Fish. Blood. % Feb. 5 if Kiel Harbour. —1:093°% 2°033 Gadus morrhua. —0-720 ” 6 II ” ” a ” ” ”» —0°75 —0:-751 pm 2 Tit 6 Ap ‘5 a Pleuronectes platessa. —0-66 (3 species.) Bp IV 3 4 oe 55 Pleuronectes platessa. | —0-65 (3 species.) oy LUG Vv On §.8. ‘ Poseidon ”’ —1:3 2:6% G. morrhua —0:758 in Baltic just out- —0:710 side Kiel Forde. —0-730 ” ” a3) P. platessa. (3species | —0:718 for each det.) —0-720 op ale VI Kattegat. —1:665 2°97 G. morrhua. —0:715 ” op as P. platessa. —0:73 Raia radiata. —1-51 mo les VII Kattegat, near Coast —1-71 3°15 G. morrhua. —0°8 of Sweden. ; —0) 77 R. batis. —1-82 March 5 XIV North Sea. —1-90 8°485 G. morrhua. —0-73 —0:79 —0:75 ; —0:77 XV Helgoland. —1-90 nS P. platessa. —0-78 —0-84 —0:75 —0:°77 XVI Helgoland. —1-90 G. morrhua. —0:778 —0:748
The “Poseidon” was fitted with a special laboratory for the examination of the
wet and live fish as they came from the trawl. investigation were all in excellent and uninjured condition.
The specimens used for the The freezing point
determinations were made at once, irrespective of whether it was daytime or
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2a.m. Since the expedition left Kiel in February, it may well be imagined that the voyage was not particularly comfortable.
As the “Poseidon” was engaged in making a complete series of scientific determinations of the oceanographical conditions prevailing, it was possible to obtain the results of chemical analyses of the sea-water at the place where the fish were captured (and within a few hours of the trawling). The freezing point of the bottom water was also taken by myself whilst the trawl was being dragged along the bottom.
A very considerable number of Plewronectes platessa were examined and the figures show a continuous but slight change from Kiel (A 0-655) to Helgoland (average A 0-787) the water having changed from /A 1:093 to A 1:90. But in the case of the cod fish the variation between individuals at one place is often greater than the difference between specimens from two such different waters as that in the Baltic and that at Helgoland.
Other species of teleosts inhabiting sea-water and brackish estuarine water, including the eel from fresh-water and from sea-water, were examined later and shown to possess a slightly higher osmotic pressure in sea-water than in fresh- water.
These results have been generally confirmed. I think we can say quite definitely today that the teleost fishes have practically achieved independence of the external medium although not completely freed from its influence. It is also clear that each different species of teleost fish has its own mean osmotic pressure, or, to put it another way, its own characteristic chemical composition (even those species living together in water of the same salinity differ slightly), which is quite in accordance with modern physiological discovery.
Modern Views on the Internal Media and their Regulation in Elasmobranch Fishes.
More exact determinations of the composition of the blood of these animals, whose strange physiological condition separates them as far from other fishes as their morphology, have revealed the following facts:
Under normal conditions in the open sea the blood of elasmobranch fishes has approximately the same osmotic pressure as that of the water in which they are living. Duval has pointed out, however, that there is not complete isotonicity. The freezing point of the blood of these fishes examined during his experiments and also, he notes, as indicated in results of some earlier workers, was more frequently a little lower than that of the surrounding sea-water.
Thus: A of blood. A of local sea-water Scyllium catulus Hie Ao 2-18 Sea-water, 2-13 (Frédéricq, 1901). Rays at Arcachon .. ts 1-89 Sea-water, 1-84 (Duval). Scyllium catulus os ie 2-17 Sea-water, 2:08 (Duval, Monaco).
The difference is small and was not considered worthy of note by the earlier workers. — 2
The low freezing point of elasmobranch blood is due to salts plus an unusual amount of urea. Actually less than half of the osmotic pressure is due to salts. Duval gives the NaCl of the serum as only sufficient to produce a A of 0:97° where the blood freezes at — 2:17. In other words the urea is responsible for a depression of the freezing point of 1-20° C. Urea may be present up to a propor- tion as great as 3%. This is extraordinary when one remembers that only 0:03%
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is usually found in mammalian blood plasma.* But this is not the only startling feature. It is not difficult to demonstrate (Dakin and Edmonds, 1931) that urea in solution diffuses very easily through living bounding membranes of aquatic animals, and urea is one of the most diffusible substances through artificial membranes. But the gill membranes of the elasmobranchs must be impermeable to urea. And then, notwithstanding this, one finds (Duval, 1925) that the red corpuscles are totally unaffected osmotically by the urea in the blood. The corpuscular wall would appear to be easily permeable. Truly a paradoxical state of affairs.
It is no wonder that considerable interest has been paid to the effect on the elasmobranchs of altering the constitution of the sea-water in which they are held. Unfortunately sharks and rays are almost entirely confined to sea-water, and it was a long time before any from natural fresh-waters were examined. It was quite easy to see that in experimental tanks the osmotic pressure of the blood of elasmobranchs was very easily affected by changes in the salinity of the water and thus they were entirely different from teleosts in this respect.
My own figures for these fish from sea-water in the Baltic and North Seas showed that:
Raia radiata, in the Kattegat, with A of sea-water 1:66, gave blood A 1:5, whilst Raia valonia, in the North Sea, with A of sea-water 1:98, gave blood A 2:0.
Duval showed that when a dogfish was put without gradual change into a sea-water diluted so that its freezing point was only —1:07 (the normal was —2:08° C. where he was working), the freezing point of the blood changed from —2:17° C. to —1:76° in 3 hours 30 minutes.
Other experiments have shown that the normal close agreement between the osmotic pressure of elasmobranch blood and that of the sea-water is only found within certain restricted limits. It would appear then that the elasmobranch is not entirely without control over its internal media after all. All these experi- mental changes of salinity result, however, in serious damage to the fish, and it seems peculiarly important in this work that conclusions are not drawn from unhealthy specimens. Certainly Duval’s experiments lasted for too short a time and were accompanied by too abrupt and deleterious a change of medium to indicate whether a real and new equilibrium had been arrived at between the fish and its environment.;
In 1931, H. W. and C. G. Smith solved. the problem of the conditions prevailing in elasmobranchs by making a journey to Siam and Malaya where certain species were to be obtained swimming in perfectly fresh water.
The osmotic pressure of the blood of these typically marine fish in fresh-water corresponded to a A of 1:0° C. It is obvious, therefore, at a glance, that the elasmobranchs are no more incapable of upholding the osmotic pressure of their internal media than are the teleosts or fresh-water crustacea. Their degree of independence may be different, the physiological mechanisms involved may be different, but it is a clear fact that in fresh-water the highest aquatic inver- tebrates, and the aquatic vertebrates all sustain internal media (the blood is
* Baglioni found about 2-61% in elasmobranchs at Naples and showed that to sustain the normal beat of the heart of a shark an artificial saline solution had to contain 2 grammes urea and 2 grammes of NaCl for every 100 c.c. of water.
y I gather that R. Margaria still considered in 1931 that the elasmobranchs were unable to sustain a difference between the body fluids and the environment. This was again the result of experimental methods on a few animals.
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that particularly dealt with) whose salinity is controlled and which is in most cases actually higher than that in the blood plasma of terrestrial mammals.
Smith’s results indicated the probable solution of another problem which has interested me for many years. The fall in osmotic pressure in the fresh-water elasmobranchs could conceivably be due to the mere absorption of water by the fish—the gills or other bounding membranes acting as an impermeable membrane to salts and urea but permeable to water. By analysis of both the urea and the chloride contents, Smith has shown, however, that the concentration of these substances is not reduced equally during the passage to fresh-water. There is a big fall in the urea concentration (a fall of 70%), but a fall of only 25% in the chlorides.
This would indicate that the regulation is much less simple than might be supposed, and that in regard to the salinity of the blood the elasmobranch is not acting very differently (if it is different at all) from the teleost fish. The part played by the urea in the physiology of the elasmobranch is the peculiar feature— in fact it is unique in the animal kingdom.
Before we turn to the significance of these facts concerning the aquatic vertebrates of the two groups, elasmobranchs and teleosts, let us consider in greater detail the results of modern researches on the aquatic invertebrates.
The Relation between the External Medium and the Internal Medium of Marine and Fresh-water Invertebrates.
The earlier researches showed, as I have already pointed out, that both the salinity and osmotic pressure of marine invertebrates were very like those of their surroundings. But it was very soon observed that some species of aquatic invertebrates, which lived in fresh-water and belonged to the same animal groups as the typical marine forms, managed to conserve a high salinity for the blood and a high osmotic pressure, although immersed in fresh-water. This resulted in a tendency to divide the aquatic invertebrates into two sub-divisions, the marine and the fresh-water forms, and to assume that a very different physiological function had been evolved in the latter. Bottazzi himself (1908) introduced the term “poikilosmoticity” for the marine invertebrates, assuming that their salinities fluctuated (and with an isotonicity) with that of the surroundings, like the temperature of “cold blooded” or poikilothermic animals. The fact, known at that time, that the blood of a marine crab placed in almost fresh-water, or even in a mixture 50% sea-water and 50% fresh-water, never reached isotonicity with its surrounding medium, was regarded as due to the inability of the crab to live long enough in the diluted sea-water for the state to be attained.
The facts are as follows: In most marine invertebrates, when living under ocean or open sea conditions such that the sea-water is of a salinity between 33% and 35%., the osmotic pressure of the body fluids is approximately the same as that of the sea-water. (There are, however, some interesting discrepancies even here.) When, however, the salinity is considerably reduced, either by the addition of water in experimental tanks or where estuarine and river conditions arise, the osmotic pressure and salinity of the blood and other fluids both fall until a new equilibrium is reached, but whether this results in a new isotonicity depends entirely upon the amount of dilution of the sea-water and upon the species of animal concerned. In no case does the internal fluid become isotonic with the external if the sea-water is very considerably diluted. The closest
approach to this condition is seen in animals such as worms and molluscs with Cc
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extensive unarmoured body walls. (1 always assume that the creatures remain alive and reasonably healthy.)
The crab, Heloecius cordiformis, which is found in certain estuaries of New South Wales, presents a freezing point for the blood a little below that of the sea- water at A 1:9, in which the animal is living. When, however, the animal is placed in fresh-water, the salinity of the blood changes as indicated in the following table (Dakin and Hdmonds, 1931).
Reaction of the Crab, Heloecius cordiformis, to fresh-water.
Duration of : A of Medium. A of Blood. Experiment. Medium. (° C.) (@ @) 2 hours Fresh-water. 0:0 1-9 (anions “ i 0:0 1:83 Sites . i 0-0 1:7 Wh hg Ps 0-0 1-56 BO) 35 Diluted sea-water. 0-1 1-3 8 weeks 55 6p a 0:72 1°38 Controls in sea-water. 1:98 1:89
But the rate of change indicated here varies for different invertebrate species and many marine invertebrates cannot withstand the conditions of the above series of experiments at all, but could only be tested in less diluted sea-water.
Schlieper (1930) has recorded that the crab, Carcinus maenas, which also lives under a wide range of conditions on the European Coast, presents at Helgoland a freezing point for the blood approximately the same as that of the sea-water there (A 1-9 or thereabouts), but in the Baltic Sea, where the freezing point is only 0-75° C. lower than that of fresh-water, the A for the blood is retained at 1:48° C. to 1:55° C. So Carcinus maenas is evidently very similar in its reactions to our Heloecius. ,
Contrasted, however, with the above types are other exclusively marine crabs, such as Portunus puber, Herbstia condyliata, and Maja verrucosa, whose blood freezing points are normally exactly the same as the sea-water in which they are living. Schwabe’s experiments (1933) show that when Maja verrucosa is placed in diluted sea-water A 1:33, the blood has exactly the same A after 36 hours as the diluted sea-water.
One of the most surprising examples is that of the two species of Nereis, Nereis pelagica and Nereis diversicolor. Schlieper’s experiments (1929) showed that the latter species, when placed out of sea-water of 32%, into that of 15%, seemed quite normal after 24 hours, whilst the former swelled up through osmotic intake of water and died. WN. diversicolor is apparently able to sustain an osmotic pressure for its body fluids greater than that of the environment when the latter is brackish water (A 0:21° C.), the other species is not.
The marine worm, Arenicola marina, whose internal medium has a A 1:70 in a sea-water of A 1:72 (in the North Sea), has a A of as low as 0-75 for its internal medium in the Baltic Sea water at 0:77. This A is lower than that of many aquatic crustacea in the perfectly fresh water of rivers and lakes. For example, the A for the blood of the fresh-water crayfish is 0-8 to 1:0° C., and for the fresh-water crab, Telphusa fluviatilis, 1:16° C. We (Dakin and Edmonds (1931)
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and Edmonds (unpublished) ) have shown that in a _ salt-water mollusc (Onchidium) and in five other species of Crustacea, the osmotic pressure of the blood in diluted sea-water is sustained in defiance of the external medium (i.e., the creatures are homoiosmotic).
This practically sums up the whole position so far as a change towards lowered salinity is concerned. The invertebrates in fresh-water represent then the species which can withstand an outer medium which is deficient in salts. Of these, the higher crustacea, like the crayfish, present a freezing point for the blood of —0:8° C. to —1:0° C., and so are not unlike the marine crab, Heloecius cordiformis, referred to above, and to other homoiosmotic forms, except that the modification of the external environment has been greater. In contrast with these are the fresh-water lamellibranchs, which present the lowest osmotic pressure for their blood and internal media. Thus the pond mussel, Anodonta, gives A 0-1 in fresh-water. Even here, however, after thousands of generations of fresh-water existence, the animal sustains an internal fluid with a higher salinity than that of the surrounding medium.
Poikilosmoticity in Marine Invertebrates.
We have taken for granted that within some limits, which vary for different species, the osmotic pressure of the internal media of marine invertebrates is almost identical with that of the surrounding sea-water. More accurate experi- ments in which a large number of individual species are used may show that even this is not so general as has been supposed.
Schlieper consistently obtained a A for the blood of Carcinus maenas which averaged 1:96° C. when the external sea-water was —1:91° C. Duval also noticed a frequent slight hypertonicity. Other workers have observed the same thing. In Pachygrapsus crassipes, however, the blood only freezes at —1:327° where the sea-water freezes at —1:975° C. (Baumberger and Olmstedt, 1928).
Miss Edmonds has made a special study of these conditions in the New South Wales crab, Heloecius cordiformis, and here there seems to be a regular hypo-tonicity, the difference in the freezing points of sea-water and blood being about 0:25° C. She also found similar conditions for Leptograpsus variegatus.
It is a curious and important fact that, for certain marine invertebrates which have been the subject of experiment (not so many have been used in this way), an increase in the concentration of the salinity of the external medium produces a greater and more speedy effect upon the internal media than does a dilution. Thus, according to Frédéricq and Duval, when Carcinus maenas is immersed in sea-water which has been concentrated, the salinity of the blood rises after a few hours in apparent exact correspondence with that of the external medium. Schwabe (1933) found that the three marine crabs, Dromia vulgaris, Herbstia condyliata and Portunus corrugatus, attained almost isotonicity in 51%, sea-water after only 48 hours.
Our experiments with the brackish-water crabs of New South Wales are interesting in this respect. In an early paper (Dakin and Edmonds, 1931) we found that whilst an increase in salinity in the blood of Heloecius cordiformis took place in concentrated water, the freezing point of blood was only — 2:92 in water of freezing point —3-28 after 28 days. Further experiments of Edmonds showed that in salt solutions of A 3:24, crabs, after a duration of 36 days, presented a freezing point for the blood which was only 0-14° C. different from that of the sea-water. Heloecius agrees, therefore, with Carcinus and other species in its
xX PRESIDENTIAL ADDRESS.
reaction to the highly concentrated saline medium, except that it certainly “gives way” more slowly to the influence of the environment.
I am not altogether in agreement with Schlieper when he says that in contrast to all fresh-water animals and marine teleosts which have mechanisms for controlling their water content, most marine invertebrates are poikilosmotic and have an osmotic pressure which is the same as that of the external medium. The latter may be the case, though only within certain limits, but is it correct to say that the fresh-water species have a mechanism not possessed by their marine relatives? Is it correct to assume that there is some new mechanism at work in Nereis diversicolor which is not present in its related species, N. pelagica?
The real poikilosmotic condition often only exists (if it does show itself) between narrow limits. It may be quite true to say that a certain species of marine crab is poikilosmotic in sea-water between certain limits of salinity, but the use of the term is unfortunate if it means that aquatic invertebrates are to be divided into two sharply-marked classes, poikilosmotic and homoiosmotic. The differences between distinct zoological groups like the echinoderms, with their vast coelomic cavity, and crustacea, with a haemocoele, is another matter. The important difference between species which can invade and live in brackish or fresh waters and their relations limited to the sea, is that the former are capable of tolerating a change in the constitution of their internal fluids and of keeping in action a series of processes which sustain new equilibrium and a “steady state”. The problem may be wholly quantitative, if I can put it this way, rather than qualitative.
This is a convenient place to refer to a lesser known field, to the conditions which obtain in typically aquatic invertebrates, such as the crabs (and some other crustacea), which have invaded the land and which live a more or less terrestrial existence. .
Observations of A. S. Pearse (1932) show that there is quite a range of types. In Gecarcinus littoralis which lives in burrows, often at considerable distances from the sea, the A of the blood is only 1:65, whereas the nearby ocean water has a A of 2:04. Cardisoma guanhuni, another very large crab, often found far from the sea, has a similar freezing point for the blood.
Pearse concludes that land crabs have blood of lower osmotic pressure than those of marine crustaceans and that the attainment of land life (possibly through the acquirement of air-breathing habits) is associated with a reduction in the salinity of the blood.
So far we have considered the osmotic pressure of the internal media and assumed, perhaps more particularly in the case of the marine invertebrata, that it was largely due to sodium chloride, together with the other salts found in sea- water. This was the thesis of Quinton (1897), referred to at the beginning of this address. And it is true that in the relative amounts of the inorganic constituents of the body fluids even of the higher animals—land animals and also fresh-water animals—the general resemblance to the composition of sea-water is most striking. ;
In the case of the marine invertebrates, an absolute identity of the saline constituents of the body fluids with the proportions of the salts in the surrounding water has been too often assumed just as an exact isotonicity and a complete dependence was taken almost as universal after the early work on this subject.
Such is, however, not by any means the case, although the divergences may be small.
PRESIDENTIAL ADDRESS. xxi The following table from Pantin will serve to set the matter out. I have added the last lines giving the figures for the teleosts and for human serum. Date Taken or Calculated from. Na. Ke Ca. Mg. (OL SO,. [XB Sea-water Dittmar (1884) 100 3°6 3°9 12) 181 20:9 ee 53 2 ae — — — — — — —2-0-2:-4 Aurelia flavidula (mesogloea) Macallum (1926) 100 52 4-1 11-4 186 13-2 — Limulus polyphemus M5 is 100 5:6 4-1 11-2 187 13-4 —2:04 Aplysia limacina Bethe (1929) 100 4-0 4-4 11 180 — = Ms nA -- | Quagliariello (1925) | — = _ 2-32 Homarus americanus | Macallum (1926) 100 3°7 4:9 1O7/ 171 6:7 = Acanthias vulgaris 5 5S 100 4°6 2°7 2°5 166 — —2:04 Carcinus maenas Bethe (1929) 100 4:8 4-5 4-8 180 —_— Variable. Frog Macallum (1926) 100 | 11:8 3:17 0-79 135-6 _— —0°4 Dog a3 Xe ve PA 100 6-6 2-8 0-76 139-5 — —0°6 “Hard ”’ fresh-water (Wembury) .. | Pantin (1931a) 100 | 74 299 66 190 95 = (Cod) Gadus callarius | Macallum (1926) 100 9-5 3°93 1°41 149-7 — = (Pollock) Pollachius virens ie 45 100 4-33 3°10 1:46 137°8 = = Human serum Kramer and Tisdall (1922) > |} 20) 5:97 2-99 0-896 | 106-46 5-73 — Human serum (Another analysis) 100 6°75 3:10 0-69 128-8 — =
It is clearly evident from an examination of the few cases in this table that there is a range of dependence upon the saline composition of the sea-water in marine invertebrates, just as there is in osmotic pressure. Thus, the body fluids of the molluscs, as exemplified by Aplysia, are in close agreement with sea-water, whilst the higher crustacea, as exemplified by the lobster (Homarus americanus) and the crab (Carcinus maenas), agree pretty closely in the relative proportions of sodium, potassium, calcium and chlorine, but are very different on the score of magnesium. Unfortunately, there are too few complete analyses available of the media of invertebrates. The vertebrate animals are still more different but agree very closely amongst themselves.
These facts are very significant because, as will be pointed out later, there is evidence now to show that the bounding membranes are not impermeable to ions, although they are probably more permeable to water.
It is extremely probable, indeed one might say certain, that any control of the salinity and osmotic pressure of the internal media of aquatic animals is not effected merely by changes in water content. The body fluids are not merely a diluted or concentrated external medium. Their ionic composition is a function of the phenomena of protoplasm itself and of the body as a whole. The factors behind these phenomena are very elusive and the interpretations of the facts may well involve not only a physiological investigation but a palaeontological study.
The Evolutionary Origin of the Independence of the Body Fluids of the Vertebrates. Before proceeding to the final stage in this subject, which is at the same time the most puzzling and the most interesting—I mean the search for the
XXii PRESIDENTIAL ADDRESS.
-mechanism and sources of energy which enable a creature to sustain internal media of a certain salinity in defiance of the composition of that which bathes its delicate bounding membranes—it is desirable to return to that historical problem raised 46 years ago by Bunge (1889) and then again by Quinton in 1897.
Quinton, I may remind you, stated that, so far as salinity was concerned, the blood of most animals was an altered sea-water. In 1903 the Canadian physiologist Macallum, apparently unaware of the suggestions of Bunge or Quinton, advanced the view that the blood plasma of vertebrates and invertebrates with a closed circulatory system is, in its inorganic salts, but a reproduction of the sea-water of the remote geological period in which the prototypic representatives of such animal forms first made their appearance.
This fascinating theory, as is often the case, caught the imagination, and today we find medical textbooks devoting some pages to the matter. In 1912, when I first criticized Macallum’s theory, I pointed out that it might be a very reasonable assumption to regard the saline composition of the body fluids of animals as a relic of early biological history. In fact, I was prepared to accept one of his statements as it stood, viz.: “the inorganic composition of the blood plasma is an heirloom of life in the primeval ocean”. But that was on the under- standing that the heirloom could be modified as it was passed on in the course of evolution and not handed on unaltered like a piece of family plate. I refused, however, to accept Macallum’s main thesis—that the blood plasma in any animals represented, so far as its inorganic composition was concerned, the composition of the sea-water of some remote geological epoch. For example, I saw no reason why because the A for teleost blood was approximately —0-6 one had to assume that the bony fishes (and, indeed, the ancestors of the higher vertebrates) had evolved in brackish water which had.a salinity corresponding to this.
The work of recent years has confirmed my belief in this matter.
Take, for example, the fresh-water crabs of certain coastal creeks flowing into the Hawkesbury River near Sydney. These crabs (still an undescribed species) present every indication of a migration into the waters where they are now found from the brackish waters of the estuary, and from the geological evidence alone one may reasonably assume that the migration has been relatively recent. The crabs are thoroughly adapted to fresh-water and the osmotic pressure of the blood corresponds to A 1:23, a figure which is also approximately that of some fresh-water crayfish.
It is difficult to see how, in either or both cases, the blood salinity represents that of the ocean sea-water from which they or their ancestors came, nor can it represent the salinity of any particular stage on the way.
The purely marine crab, Pachygrapsus crassipes, has been found to have a A for the blood as low as 1:327 where the sea-water was A 1:975° C. There is no reason to assume that this species of Pachygrapsus has evolved some new mechanism for regulating its body fluids, or that it did so in water of low salinity. The fact that a marine crab is able to sustain such a difference of composition as this between its body fluids and the external medium is additional evidence against the necessity for assuming that the ancestors of the vertebrates had their origin in an ocean of only half the present salinity or less, because the body fluids of the vertebrates of today present salinities of that order.
Take again, for example, the New South Wales estuarine crab, Heloecius cordiformis. We have found this crab particularly common where the A of the sea-water was 1:98° C. (That is to say, on flats where the sea-water is not
PRESIDENTIAL ADDRESS. XXili
diluted to any great extent except during the rains.) It also extends into regions where the water is much less saline, but not into fresh-water. (Hdmonds, unpublished paper, found it in water with a salinity corresponding to a A of 0-8 to 0:58° C.) Yet in water of the lowest salinity the A for the blood is 1:43° C., and at every point between its most saline and its least saline habitat the blood A is a function of its environment. Edmonds has placed Heloecius cordiformis taken from water of high natural salinity (A 1:98) into water with a freezing point of —0-72 and kept them in aquaria for two months. During the early part of this period the A of the blood fell to 1-:38° C., but then remained definitely constant. It was clearly evident that a new equilibrium had been reached and sustained with a very striking difference between the body fluids and the external medium, and the experimental result is practically identical with discoveries in the field. I see no reason to assume that Heloecius cordiformis evolved this brackish water homosmoticity at some particular period in the remote past when the water in which it was living was of some special salinity. The salinity of the blood of fresh-water crustacea tells nothing definite about the exact concentration of early ocean waters, although I am not prepared to argue that the duration of evolutionary existence in fresh-waters is not without effect on the saline composition of the body fluids. At the same time, even in this respect it is necessary to remember that there is no reason why the particular salinity should not be just as much a reflection of the particular physiology of the species as a reflection of anything else. Where a large number of deter- minations are made it will be found that there are considerable differences between the osmotic pressures of one crab and another even under the same conditions and in the same locality (Hdmonds).
It might be urged that the cases utilized above for this discussion are estuarine invertebrates. I would answer that I see no evidence why real marine species should not be capable of migrating into brackish waters and fresh waters today, and in such cases the salinity of the body fluids would no more represent the present day salinity of the ocean than does that of Heloecius cordiformis. But suppose we turn to the aquatic vertebrates—both the teleost fishes and the elasmobranchs (although so different physiologically) exercise a regulation over the salinity of the blood. In both cases it is almost independent of the water bathing their bodies. Actually the salinity of elasmobranch blood is some- what greater than that of teleost blood. It might be inferred from this that the elasmobranchs evolved in ocean water of a later date (Macallum utilizes very largely the calculations of Joly on the age of the earth by estimating the increasing salinity of the ocean). This, however, would be rather contrary to the usual views on the evolution of the vertebrates. Macallum’s view is that the elasmo- branchs evolved their fixed salinity at an earlier epoch than the teleosts and that it is now higher in the former because they have been exposed to the ocean’s increasing salinity for a few more million years than the teleosts. To my mind this argument, which can be used either way, weakens his thesis still further.
The aquatic vertebrates—the teleosts and elasmobranchs—which have evolved an independence of their body fluids may:
(1) Have evolved this independence in and when the ocean water had a salinity corresponding to a A of say 0:6° C. (a long time ago), and then whilst the ocean water has slowly increased its salinity to 35%, the original or nearly original salinity of the ocean has been retained in these vertebrates. (Macallum’s view.)
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(2) Have evolved in ocean water of higher or lower salinity (to any degree) and for some reason gradually fixed their salinity at the present prevailing figures.
(3) Have evolved from proto-vertebrates or early vertebrates which migrated from the sea into fresh or nearly fresh water and in which species, exactly as with the higher invertebrates which do this today, the salinity of the body fluids fell to a new equilibrium but was sustained at this, and by the consumption of energy was kept nearly independent of the vagaries of the external media.
The experimental evidence all seems to point to the latter, and it is interesting to note that many modern palaeontologists favour the view that the vertebrates were evolved in fresh-waters. (Chamberlain 1920, Grabau 1913, and O’Connell 1916, and others. Marshall and Smith (1930) affirm that the vertebrate glomerular kidney must have evolved in fresh-water.)
The Mechanism whereby the Steady State is Maintained.
It may be well to point out that recent researches have shown only too clearly that the mechanism whereby aquatic creatures sustain body fluids markedly independent of the external watery environment is by no means as simple as was once supposed.
The early workers spoke of “closed” blood systems and impervious body walls. Macallum regarded the kidneys as the essential regulators, and even in his paper of 1926 holds to this view. Thus: “The low concentration of salts in the blood, as compared with the concentration of salts in sea-water, and the maintenance of the palaeo-ratios in Selachians, after very many millions, possibly hundreds of millions of years of life in the sea, indicate unmistakably how inflexibly constant, practically, is the action of the organ concerned, the kidney in the vertebrates.” On another page he says: “There are in Invertebrates no structure or structures having a function or functions quite similar to those of the vertebrate kidney... .” | “In the long ages the kidney has ever thus performed functions which, for constancy and regularity, are unrivalled in the world of life, except by those of the cell nucleus, which, of course, is of vastly more remote origin. This constancy contrasts with the variations in functions which the other organs in vertebrates have undergone. It has made the vertebrates, with all their ranges of development, possible. Without such a constancy there could be no change in habitat from sea to land and fresh-water and back again to sea, for with such a change there would be a variation in the inorganic composition of the internal medium, an impossible handicap in the struggle for existence, which would greatly affect the development of the organs after the EHo-vertebrate stage was passed.” But Macallum believed that the live bounding membranes of aquatic animals are impermeable to saits. Apart from this, his views do not conform to the facts.
So far back as 1910 it was shown by me that the osmotic pressure and salinity of the contents of certain marine teleost fish eggs were quite unlike that of the sea-water in which the eggs were floating, but only so long as the egg membranes remained alive. At the same time the egg of the elasmobranch was shown to have a low freezing point (1:80° C.) similar to the blood of the adult fishes of the same group, and in 1928 Needham and Needham recorded 888 mg. of urea in such eggs. There was distinct evidence in both cases of the action of the bounding membranes. And again, when discussing Macallum’s views (Dakin, 1912) it was stated that “The bounding membranes of the body and the fluids
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bathing them are the prime factors in the regulation of the blood constitution so far as salinity is concerned”. “It may be said that for the substances for which it is permeable it (the bounding membrane) does not behave as a dead parchment membrane; on the other hand it exerts a direct and powerful influence.’
Modern research has fully borne out these conclusions. The researches of Smith (1930), confirmed by Keys (1931) and Bateman and Keys (1932), have shown for example that the teleosts swallow sea-water and actually secrete chloride by the gills. Keys (1933) regards the facts at present as proving that in teleosts the kidneys conserve the saline constituents and eliminate water, whilst excess salt is eliminated by the gills, water being conserved by the bounding membranes of the latter organs.
If the regulation of the blood of the teleost fishes be controlled by the two sets of organs, we are still left very much in the dark as to how the separating membranes carry on the work. Are we to conclude that the fresh-water teleosts are different structurally from their marine relatives in so far as their kidneys are concerned? It has been suggested that these organs in fresh-water fishes are capable of very efficient water filtration combined with salt conservating powers.
It is not at all easy to devise experiments which will enable one to discover how the bounding membranes of invertebrates are functioning, but Adolph (1926) has found that frog’s skin is more amenable. Adolph’s important work has shown that whilst the frog regulates the general water content of its body by the kidneys, the inflow through its skin when immersed in fresh-water is quite definitely under control. He has shown that this inflow is due in large part to forces other than osmotic pressure. It is striking in this respect that if the skin be removed, the body wall is no longer able to function in this way. ‘‘The skinless frog is an ideal osmometer.” The real forces at work in determining and controlling the inflow are, however, stated by Adolph to be still unknown, but they are wholly in the skin.
In passing, reference should be made to a very interesting point mentioned by Adolph, which seems to me to be well worthy of attention in connection with our invertebrate findings. After showing that the exchanges of water are caused in large part by forces other than osmotic pressure, he adds that “only in the higher concentrations, where the medium is more highly concentrated than the frog’s blood and lymph, do the rates of exchange of water give any appearance of being proportional to concentration”. Is it mere coincidence that in the crabs, Carcinus, Heloecius, and the other species previously mentioned, the unknown regulating processes do not seem to function when the animals are placed in concentrated sea-water?
The work of Bethe (1929) has already been referred to. It will be remembered that the early workers believed the bounding membranes of aquatic animals to be semi-permeable (permeable to water but impermeable to salts). Bottazzi and Enriques (1901) supported this view and many others have since then taken this position (including Macallum).
In favour of this attitude is the fact that many marine invertebrates, when placed in diluted sea-water, swell up and increase in weight. The bounding membranes act like the semi-permeable membrane of an osmometer. A starfish shows this particularly well. But it can be seen equally well at first in worms and other types. If, however, the experiment can be withstood by the animal concerned and the duration is not just that of a few hours, it will be found that in many cases the initial increase in weight disappears. The explanation
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is that the membranes were more readily permeable to water than to salts, not that they were impermeable to the latter.
As Schlieper points out, however, there is room for further experiments along these lines.
Our own work (Dakin and Hdmonds, 1931) confirms that of Bethe and others in showing that in many aquatic invertebrates (including the Crustacea) the bounding membranes are permeable to both water and salts. Yet the body fluids remain constant in a particular environment. This applies not only for species like the Crustacea, in which a protective impermeable body wall has been evolved except for areas like gills, but for types such as Oligochaete worms, with their soft dermo-muscular body walls.
If the external environment is changed, the body fluids change too—but not to the same extent—a new equilibrium is reached and once again a steady state is attained. 3
But how is the steady state sustained?
Schlieper has shown that, in some cases at least, an extra consumption of oxygen is required to provide the energy, and that this can be experimentally demonstrated. Thus in the crab Carcinus the need for oxygen increases with any diminution in salts in the external medium. Curiously enough, however, another crab (Hriocheir) presents no such increase in its energy requirements on passing from salt- into fresh-water.
Schlieper concludes that the excretory organs are not concerned in the osmotic control of the blood of Hriocheir and the crab Telphusa because the urine is isotonic with the blood. The same thing applies to the crab Carcinus maenas.
But the urine of the fresh-water crayfish (Potamobius) has extremely low salinity, and the A is only 0:16° when the blood A is 0:8 (Schlieper).
We are indeed far from the last word in connection with the aquatic invertebrates. There seem to be the most unexpected differences between them. If Carcinus and Heloecius are taken from sea-water and placed in fresh-water, there is no increase in weight which would result from the passage of water into the animals and yet, as we have seen, their salt content changes—Cl ions migrate outwards. But if exactly the same experiment is tried with the crab Maja verrucosa the weight increases—water is absorbed. Is it possible that in the same group of animals the organs are different in structure and function, or are the observed effects due to experimental conditions, to lack of acclimatiza- tion, etc? More than one investigator has discovered that damaged or dying gill and other membranes are permeable, whereas normally they are not so.
Pantin (1931) has investigated experimentally a peculiarly interesting example amongst aquatic invertebrates—the estuarine flatworm, Gunda ulvae. The body wall of this worm is permeable to both water and salts, yet the internal fluids are under some control and the animal will withstand considerable change in salinity. In fact, for some reason, it seems to prefer a changing medium such as one meets in a tidal estuary. If placed in fresh-water completely minus salts (i.e., distilled water), the worm swells greatly and dies; but if a small quantity of calcium be added the animal can survive much longer.
It has long been known, of course, that calcium ions have very definite effects upon the permeability of protoplasmic membranes,* and it is very well
*R. K. 8S. Lim showed in 1917 that Carcinus maenas, the crab so often referred to in this paper, lived longest in fresh-water when calcium was added, and stated that this was due to altered permeability of the membranes.
PRESIDENTIAL ADDRESS. XXxvii
known that a solution of common salt of the concentration of sea-water is likely to be as poisonous to marine animals as fresh-water. The addition of calcium ions seems to antagonize the sodium ions. The exact proportion of the different ions is indeed a matter of considerable importance in setting up aquaria with different salt solutions. And it needs only a few experiments to realize that marine animals which are never found in brackish water will live for a time in highly diluted sea-water when fresh-water proves almost immediately fatal.
Finally, it is a most important fact that the saline independence of the body fluids of the aquatic metazoa is not dissimilar from the conditions found in the cell itself.
The study of the live bounding membranes of the aquatic animals is really only just beginning. Two recent discoveries may serve to illustrate this point. Schwabe (1933), for example, has shown that in the crabs Carcinus and Eriocheir, when ecdysis is taking place, there is a fall in the concentration of the blood and the crabs swell up owing to intake of water. It seems clear that this is due to changes in the body wall.
Again the famous French physiologist Paul Bert made the discovery that eels which had been carelessly handled so that the mucus (so characteristic of the skin) had been removed, were no longer able to withstand a sudden change from fresh- water to sea-water. Duval took the matter up again and conducted a very interesting series of tests to see if it were true, and if it were due to partial loss of control over the salinity of the blood. He found that, whereas the A of the blood of a fresh-water eel placed in sea-water of A 2-:13° C. was only 0:79° C., that of an eel deprived of its mucous covering was as great as 1:15° C. Now the secretion of mucus is very characteristic of aquatic animals, but I am not aware of any researches as to its function.
The evidence collected in this paper shows clearly how aquatic animals are varyingly dependent upon their environment. We, with our impervious skins, may easily fail to realize the sensitivity to a changed medium which may be experienced by a marine fish. It is certain that many of the migrations of aquatic animals are due to but slight changes in saline composition. In this connection the New South Wales coast provides us with excellent examples. The Peneid prawns, as you know, enter our coastal lakes as tiny larvae; they feed and grow in these enclosed waters (which have a salinity much less than the sea-water ) until they reach sexual maturity. Then they pass out to sea. It has already been shown that the freezing point of the blood of the crab Hriocheir rises after egg production. Probably this may be due to a special need for Cl during the breeding season. Schwabe (1933) believes that it is for this reason that breeding Hriocheir are never found in fresh-water. This might also be the explanation for the Peneid migration to the ocean.
It would take me too far here to enter upon this subject of migration so
far as teleost fish are concerned—so many other factors enter the field.
I should like to point out, in concluding, that the investigations to which I have devoted some considerable time in this Address do not by any means constitute a purely academic problem. Their completion may have far reaching results in physiological research. I shall be content if I have shown what a wide field is presented by the subject, and how diverse are its ramifications.
I have always been particularly interested in these problems of the internal media of aquatic animals, because they involved such a happy combination of
XXVili PRESIDENTIAL ADDRESS.
experimental laboratory and aquarium studies with the investigation of the creatures in their natural haunts.
But they have always served to make me humble—to remind me of how little we know about the functioning of living organisms. We have now collected a mass of information on this subject of osmotic pressures of body fluids. Enough to show that emancipation from the external environment was an essential preliminary to the evolutionary height realized by the birds and mammals. But the modus operandi of the regulating “mechanism” still eludes us.
Professor A. V. Hill (1931), in referring to it, summed up the position excellently when he said: “Throughout we are involved, not with general equilibrium, but with conditions maintained constant by delicate governors and by a continual expenditure of energy. How that energy is supplied, how it is utilized to maintain the structure and the organization, is, I think, the major problem of Bio-physics today.”
APPENDIX I. Freezing Points of Body Fluids °C. A=depression of freezing point below 0° C.
Internal Urine. External Species. Medium. A. “Co Medium. Author. A AO; i XGc Marine Ani|mals. Coelenterata— Aleyonium palmatum ue oe 2-195-2-196 PAO) Bottazzi. Echinodermata— f Asterias glacialis .. une Sue 2-295 2-195-2:36 He Holothuria poli ae ie Se 2-299 2-195-2-36 a Annelida— Sipunculus nudus .. 2: 20—2-31 2-29 Ff Aphrodite aculeata .. . 2-259 2:29 au Arenicola marina (Helgoland) .. iLs'7/ 72, Schlieper. 55 es (Baltic Sea) On7S 0:77 . Mollusca— Aplysia limacina Bo By, 2-195-2-360 Bottazzi. Cassis sulcosa 2-36 Di2i2, Monti. Ostrea edulis 223 2-11-2-14 te Mytilus edulis 2-26 2-11-2-14 ob Octopus vulgaris 2-16 2:11-2:14 ss Arthropoda— Limulus polyphemus 1:90 1-82 N. Rogers. Homarus vulgaris 2-29 2-269-2-278 Bottazzi. Homarus americanus 1-82 1-80 N. Rogers. Hyas aranea 1:83 1-80 Schlieper. Carcinus maenas Boil7/ 1:96-1:99 Monti. s 5 1:97 1-92 Schlieper. 3 x 1:95 1-95 1:92 38 Maja verrucosa Ae He Boils PIU? Frédéricq. Cancer pagurus aa An 5% 1:84-1:91 1-91 Dakin. Heloecius cordiformis 1:95 2:17 Edmonds (unpublished) 45 ‘3 1:38 0-72 5 50 Pachygrapsus crassipes 32 1:97 Baumberger and Olmstedt. Leptograpsus variegatus (Sydney). . 1:95 2°13 Edmonds (unpublished). Tunicata— Ascidia mentula .. sis ws 2-08 1:98 Duval and Prenant.
PRESIDENTIAL
APPENDIX I.—Continued.
ADDRESS.
Freezing Points of Body Fluids °C.
Internal Species. Medium. i Xs Elasmobranchiata— Scyllium canicula Bowe Mustellus vulgaris .. 2-36 Carcharias littorina 2-03
Trygon violacea Raja undulata ” 9” Seyllium stellare Raia radiata
» valonia », clavata Holocephali—
Callorhunchus Millit
Teleostei— Pleuronectes platessa
»” ”
3 flesus ..
” ”
Gadus morrhua
” ”
as aeglefinus Lophius piscatorius Conger vulgaris
Charanx puntacco ..
Cerna gigas
Artemia salina
RPNrN eR Fe Pb bo
“04 *034
Ani|mals from Salt-w Blood. Osmotic pressur of 1:2% NaCl of 1:3% NaCl
Fresh-water A A of Internal
Medium. Mollusca— Anodonta cygnea 0:09 Unio pictorum 0-15 Timnaea stagnalis .. 0:22-0:23 Crustacea— Daphnia magna 0- 20-0 - 67 Potamobius astacus ~ 0-80 Telphusa fluviatile .. hele Eriocheir sinensis 1-09 Potamobius astacus : Ss 0:8-1:0 Astacopsis (Australian Crayfish) 1-1 Telphusa fluviatilis 1:18
Unnamed species of crab from tributary of Hawkesbury River,
N.S.W.
1-4
Urine. I Or
2-40
Concentrat between
(Almost XS it
0:64
ater Lakes. e=to that Sol. Sol.
nimals.
Urine.
0:2
1:2
A =depression of freezing point below 0° C.
SRORAIEN:
External Medium. Author. A “Ch 2°15 Duval. 2°29 Bottazzi. 1-83 N. Rogers. 2-29 Bottazzi. 1°84 Duval. 1-88 s 2-0? Bottazzi. 1-66 Dakin. 1:9 a3 1:9 55 ion of sea-water 1-5-1-85 5 1-9 FA 1-093 3. 1:91 ag fresh-water. ot taken.) 35 1-9 Dakin. 2-0 Bottazzi. 1:92 Dakin. 1-92 3 2:14 Duval. 2-29 Bottazzi. 2-29 A Salt-water. 4-5% NaCl. Medwedewa. 8:0% NaCl. a A of External Medium. — W. Koch. — N. Monti. 0:02-0:03 Frédéricq. — Fritzsche. — Frédéricq. a Duval. — Schlieper. = Herrmann. = Dakin. — Schlieper. = Dakin.
p:0.0:4 PRESIDENTIAL ADDRESS.
APPENDIX I.—Continued. Freezing Points of Body Fluids °C. A =depression of freezing point below 0° C.
Internal Urine. External Species. Medium. A AOE Medium. Author. Se Neaes Teleostei— Barbus fluviatilis 0-50 — Frédéricq, Leuciscus dobula 0:45 = Be Cyprinus carpio 0-50 —_— Duval. Salmo fario .. 0:57 _— Dekhuyzen. Anguilla anguilla 50 0-62 — Duval. ae ae ot 56 a 0:-57-0:58 —_ Dakin. Pe 5 ica 0-61 = Keys. te (in sea-water) 0:73 abotey?/ 5 Ariabas testudineus* 0-64 _ Pearse. Ophiocephalus striatus* 0-57 — A Dipnoi— Epiceratodus (Neoceratodus) forstert 0:42 —_— Dakin. Semi-|terrestrial (Littor|al or Sea Cjoast). Crustacea— Ocypoda albicans .. ds a 1-70 2-04 Pearse. Coenobita clypeatus (land hermit crab) at Aa ae or 2-09 2-04 a5 Gecarcinus littoralis (lives some distance away from sea, on land) 1-65 2:04 35
* Air-breathing fishes from Siam.
Bibliography. !
ADOLPH, HE. F., 1925.—The Passage of Water through the Skin of the Frog and the Relations between Diffusion and Permeability. Amer. Journ. Physiol., 73, pp. 85-105.
, 1926.—The Skin and Kidneys as Regulators of the Water Content of Frogs. Amer. Journ. Physiol., 76, pp. 214-15.
—, 1930.—Living Water. Qwart. Rev. Biol., 5, pp. 51-67.
ATKINS, W. R. G., 1909.—The Osmotic Pressures of the Blood and Eggs of Birds. Scient. Proc. Roy. Soc. Dublin, 12, pp. 123-130.
BATEMAN, J. B., and Kays, A., 1932.—Chloride and Vapour Pressure Relations in the Secretory Activity of the Gills of the Eel. Journ. Physiol., 75, No. 2, pp. 226-240.
BAUMBERGER, J. P., and OumstTepT, J. M. D., 1928.—Changes in the Osmotic Pressure and Water Content of Crabs during Molt Cycle. Physiol. Zool., 1, p. 531.
BERNARD, C., 1859.—Lecons sur les Propriétés physiologiques et les Altérations pathologiques des Liquides de l’?Organisme. Tomes I and II. Paris.
, 1885.—Introduction 4 l’étude de la Médecine Experimentale. Paris.
Bert, P., 1871.—Sur les phénoménes et les causes de la mort des animax d’eau douce qui l’on plonge dans l’eau de mer. OC.R. Acad. Sci. Paris, 73, pp. 382-5, 464-7. ———., 1885.—Animaux d’eau douce dans l’eau de mer, animaux d’eau de mer dans eau dessalée, animaux d’eau de mer dans l’eau sursalée. C.R. Soc. Biol., 37,
525-27.
BetHeE, A., 1929.—Ionendurchlassigkeit der Korperflache von wirbellosen Thieren des Meeres als Ursache der Giftigkeit von seewasser abnormer Zusammensetzuneg. Pflugers Arch., 221, pp.- 344-362.
Borrazzi, F., 1897.—La Pression Osmotique du Sang des Animaux marins. Arch. Ital. de Biol., Tome 28, pp. 61-72.
, 1908.—Osmotischer Druck der einzelligen, pflanzlichen und _ tierischen Organismen. Ergeb. Physiol., Jahr. 7, Abt. i and ii, pp. 161-402.
PRESIDENTIAL ADDRESS. xXxxi
Borrazzi, F., und ENRIQUES, 1901.—Uber die Bedingungen des Osmotischen Gleichgewichts, Arch. f. Anat. wu. Physiol., Physiol. Abt., Suppl. Band I, pp. 109-170. BOUSSINGAULT, 1872.—C.R. Acad. Sci., Tome 75. BUNGE, 1889.—Lehrbuch der Phys. u. Pathol. Chemie. Leipzig. DAKIN, W. J., 1908.—The Osmotic Concentration of the Blood of Fishes taken from Sea Water of naturally varying Concentration. Bio-Chem. Journ., 3, pp. 258-278. , 1908.—Variations in the Osmotic Concentration of the Blood and Coelomic Fluids of Aquatic Animals, Caused by Changes in the External Medium. Bio- Chem. Journ., 3, pp. 478-490. ,1911.—Notes on the Biology of Fish Eggs and Larvae. Internat. Rev. der ges. Hydrobiol., Band 3. ,1912.—Aquatic Animals and their Environment. Internat. Rev. der ges. Hydrobiol., Bd. 4, pp. 538-80. , 1931.—The Osmotic Concentration of the Blood of Callorhynchus millii and Epiceratodus (Neoceratodus) forsteri. Proc. Zool. Soc. Lond., Pt. 1, pp. 11-16. ,and EHpMoNDs, ENip, 1931.—The Regulation of the Salt Contents of the Blood of Aquatic Animals and the Problem of the Permeability of the Bounding Membranes of Aquatic Invertebrates. Awst. Journ. Hup. Biol., 8, pp. 169-187. DEKHUYSEN, M. C., 1905.—Sur la pression osmotique dans le sang et dans l’urine des poissons. Arch. néerl. Sci., Ser. 2, 10. DuvaL, M., 1925.—Recherches physico-chemiques et physiologiques sur la milieu intérieur des Animaux aquatiques. Ann. Inst. Oceanog. Monaco, N.S. 2, pp. 233-407. ,et PRENANT, M., 1926.—Concentration moléculaire du milieu intérieur d’une Ascidie (Ascidia mentula). C.R. hebdom. Acad. Sci., 182, pp. 96-98. FrEpERIcQ, L., 1878.—Arch. Zool. exper., Tome 7. , 1882.—Influence du milieu extérieur sur la composition saline du sang chez quelques animaux aquatiques. Bull. Acad. Roy. Sci. Belg., 51, 8rd Ser., Tome 3, \ pp. 177-190; 38rd Ser., Tome 4, pp. 209-214. , 1885.—Influence du milieu ambiant sur la composition du sang des animaux aquatiques. Arch. Zool. exp., 2nd ser., Tome 38. , 1891.—Sur la Physiologie de la branchie. Arch. Zoologie Exper. et Gen., 2nd ser., Tome 9, pp. 117-123. , 1904.—Sur la concentration moléculaire du sang et des tissus chez les animaux aquatiques. Archiv. Biol., 20, pp. 709-730. GARREY, W. C., 1905.—The Osmotic Pressure of Sea Water and the Blood of Marine
a Animals. Biol. Bull., 8, pp. 257-270. GREENE, C. W., 1904.—Physiological Studies of the Chinook Salmon. Bull. U.S. Bur. Fish., 24, pp. 431-456. : GRIFFITHS, 1892.—Physiology of the Invertebrata. London. GUEYLARD, F., 1924.—De l’adaptation aux changements de salinité. Recherches biol. et physico-chemiques sur l’Hpinoche. Arch. Phys. Biol., 3. HaAampBurcsr, H. J., 1902.—Osmotischer Druck und Ionenlhere. Wiesbaden. Band I, II und III. ‘ HENDERSON, L. J., 1930.—Blood. Yale University Press. Hitt, A. V., 1931.—Adventures in Bio-Physics. Oxford University Press. Keys, A., 1931.—Chloride and Water Secretion and Absorption by the Gills of the Eel. Zeitsch. vergl. Physiol., 15, p. 364. ,1933.—The Mechanism of Adaptation to varying salinity in the Common Eel and the General Problem of Osmotic Regulation in Fishes. Proc. Roy. Soc., B, Vol. 112, pp. 184-199. KRIZENECKY, J., 1916.—Hin Beitrag zum studium der Bedeutung der osmotischen Verhalt- nisse ftir Organismen. Pflugers Arch., 1638, pp. 325-354. KRUKENBERG, C. F. W., 1888.—La rétention de l’urée chez les sélachiens. Ann. Mus. Hist. Nat. Marseille, 3. Lim, R. K. S., 1918.—Period of Survival of the Shore Crab (Carcinus maenas) in Distilled Water. Proc. Roy. Soc. Edin., 38, Pt. 1, No. 4, p. 14-22. Macauuum, A. B., 1904.—The Palaeochemistry of the Ocean in Relation to Animal and Vegetable Protoplasm. Trans. Canad. Inst., Vol. 7, pp. 535-562. — , 1910.—Inorganic Composition of the Blood. Proc. Roy. Soc. Lond. mn 2 aes acochemisixy of Body Fluids and Tissues. Physiol. Rev., Vol. 6,
pp. 316-357.
MarGaria, R., 1931.—The Osmotic Changes in Some Marine Animals. Proc. Roy. Soc., 107 B, pp. 606-624.
b:0:0:¢ PRESIDENTIAL ADDRESS.
APPENDIX I.—Continued. Freezing Points of Body Fluids °C, A =depression of freezing point below 0° C.
Internal Urine. External Species. Medium. IK OL Medium. Author. ky Op Is “Cie Teleostei—
Barbus fluviatilis 0-50 — Frédéricq, Leuciscus dobula 0:45 — a Cyprinus carpio 0:50 — Duval.
Salmo fario .. A 0-57 — Dekhuyzen. Anguilla anguilla .. 50 0-62 — Duval.
i 45 an he ono 0:57-0:58 — Dakin.
3 % ee 0-61 = Keys.
i (in sea-water) 0:73 1-87 % Ariabas testudineus* 0:64 — Pearse. Ophiocephalus striatus* @ok7 = 9
Dipnoi— Epiceratodus (Neoceratodus) forsteri 0:42 _ Dakin. Semi-|terrestrial (Littor|al or Sea Cloast). Crustacea— Ocypoda albicans .. oe aa 1:70 2:04 Pearse. Coenobita clypeatus (land hermit crab) aie a ais Bs 2°09 2-04 Ay Gecarcinus littoralis (lives some distance away from sea, on land) 1-65 2-04 5
* Air-breathing fishes from Siam.
Bibliography. ;
ADOLPH, E. F., 1925.—The Passage of Water through the Skin of the Frog and the Relations between Diffusion and Permeability. Amer. Journ. Physiol., 73, pp. 85-105.
,1926.—The Skin and Kidneys as Regulators of the Water Content of Frogs. Amer. Journ. Physiol., 76, pp. 214-15.
—, 1930.—Living Water. Quart. Rev. Biol., 5, pp. 51-67.
ATKINS, W. R. G., 1909.—The Osmotic Pressures of the Blood and Hggs of Birds. Scient. Proc. Roy. Soc. Dublin, 12, pp. 123-130.
BATEMAN, J. B., and Krys, A., 1932.—Chloride and Vapour Pressure Relations in the Secretory Activity of the Gills of the Eel. Journ. Physiol., 75, No. 2, pp. 226-240.
BAUMBERGER, J. P., and OLMSTEDT, J. M. D., 1928.—Changes in the Osmotic Pressure and Water Content of Crabs during Molt Cycle. Physiol. Zool., 1, p. 531.
BERNARD, C., 1859.—Lecons sur les Propriétés physiologiques et les Altérations pathologiques des Liquides de l?Organisme. Tomes I and II. Paris.
, 1885.—Introduction & l’étude de la Médecine Experimentale. Paris.
Bert, P., 1871.—Sur les phénoménes et les causes de la mort des animax d’eau douce qui l’on plonge dans l’eau de mer. C.R. Acad. Sci. Paris, 73, pp. 382-5, 464-7. ———, 1885.—Animaux d’eau douce dans l’eau de mer, animaux d’eau de mer dans eau dessalée, animaux d’eau de mer dans l’eau sursalée. OC.R. Soc. Biol., 37,
525-27.
BetHe, A., 1929.—Ionendurchlassigkeit der Korperflache von wirbellosen Thieren des Meeres als Ursache der Giftigkeit von seewasser abnormer Zusammensetzung. Pflugers Arch., 221, pp.- 344-362.
Borrazzi, F., 1897.—La Pression Osmotique du Sang des Animaux marins. Arch. Ital. de .Biol., Tome 28, pp. 61-72.
, 1908.—Osmotischer Druck der einzelligen, pflanzlichen und _ tierischen Organismen. Ergeb. Physiol., Jahr. 7, Abt. i and ii, pp. 161-402.
PRESIDENTIAL ADDRESS. Xxxi
Borrazzi, F., und ENRIQUES, 1901.—Uber die Bedingungen des Osmotischen Gleichgewichts, Arch. f. Anat. wu. Physiol., Physiol. Abt., Suppl. Band I, pp. 109-170. BOUSSINGAULT, 1872.—C.R. Acad. Sci., Tome 75. BUNGE, 1889.—Lehrbuch der Phys. u. Pathol. Chemie. Leipzig. DAKIN, W. J., 1908.—The Osmotic Concentration of the Blood of Fishes taken from Sea Water of naturally varying Concentration. Bio-Chem. Journ., 3, pp. 258-278. ,1908.—Variations in the Osmotic Concentration of the Blood and Coelomic Fluids of Aquatic Animals, Caused by Changes in the External Medium. Bio- Chem. Journ., 3, pp. 4738-490. : , 1911.—Notes on the Biology of Fish Eggs and Larvae. Internat. Rev. der ges. Hydrobiol., Band 3. ,1912.—Aquatic Animals and their Environment. Internat. Rev. der ges. Hydrobiol., Bd. 4, pp. 53-80. , 1931.—The Osmotic Concentration of the Blood of Callorhynchus millii and Epiceratodus (Neoceratodus) forsteri. Proc. Zool. Soc. Lond., Pt. 1, pp. 11-16. , and EpMONDs, ENIp, 1931.—The Regulation of the Salt Contents of the Blood of Aquatic Animals and the Problem of the Permeability of the Bounding Membranes of Aquatic Invertebrates. Aust. Journ. Eup. Biol., 8, pp. 169-187. DEKHUYSEN, M. C., 1905.—Sur la pression osmotique dans le sang et dans l’urine des poissons. Arch. néerl. Sci., Ser. 2, 10. Duvatu, M., 1925.—Recherches physico-chemiques et physiologiques sur la milieu intérieur des Animaux aquatiques. Ann. Inst. Oceanog. Monaco, N.S. 2, pp. 233-407. ,et PRENANT, M., 1926.—Concentration moléculaire du milieu intérieur d’une Ascidie (Ascidia mentula). C.R. hebdom. Acad. Sci., 182, pp. 96-98. FrEpERIc@, L., 1878.—Arch. Zool. exper., Tome 7. , 1882.—Influence du milieu extérieur sur la composition saline du sang chez quelques animaux aquatiques. Bull. Acad. Roy. Sci. Belg., 51, 3rd Ser., Tome 3, : pp. 177-190; 8rd Ser., Tome 4, pp. 209-214. , 1885.—Influence du milieu ambiant sur la composition du sang des animaux aquatiques. Arch. Zool. exp., 2nd ser., Tome 3. , 1891.—Sur la Physiologie de la branchie. Arch. Zoologie Exper. et Gen., 2nd ser., Tome 9, pp. 117-1238. , 1904.—Sur la concentration moléculaire du sang et des tissus chez les animaux aquatiques. Archiv. Biol., 20, pp. 709-730. GARREY, W. C., 1905.—The Osmotic Pressure of Sea Water and the Blood of Marine
“4 Animals. Biol. Bull., 8, pp. 257-270. GREENE, C. W., 1904.—Physiological Studies of the Chinook Salmon. Bull. U.S. Bur. Fish., 24, pp. 431-456. ; GRIFFITHS, 1892.—Physiology of the Invertebrata. London. GUEYLARD, F., 1924.—De l’adaptation aux changements de salinité. Recherches biol. et physico-chemiques sur l’Hpinoche. Arch. Phys. Biol., 3. HameBurGemr, H. J., 1902.—Osmotischer Druck und JIonenlhere. Wiesbaden. Band I, II und III. ‘ HENDERSON, L. J., 1930.—Blood. Yale University Press. Hitt, A. V., 1931.—Adventures in Bio-Physics. Oxford University Press. Keys, A., 1931.—Chloride and Water Secretion and Absorption by the Gills of the Hel. Zeitsch. vergl. Physiol., 15, p. 364. ,1933.—The Mechanism of Adaptation to varying salinity in the Common Eel and the General Problem of Osmotic Regulation in Fishes. Proc. Roy. Soc., B, Vol. 112, pp. 184-199. KRIZENECKY, J., 1916.—Hin Beitrag zum studium der Bedeutung der osmotischen Verhalt- nisse ftir Organismen. Pflugers Arch., 163, pp. 325-354. KRUKENBERG, C. F. W., 1888.—La rétention de l’urée chez les sélachiens. Ann. Mus. Hist. Nat. Marseille, 3. Lim, R. K. S., 1918.—Period of Survival of the Shore Crab (Carcinus maenas) in Distilled Water. Proc. Roy. Soc. Edin., 38, Pt. 1, No. 4, p. 14-22. Macauuum, A. B., 1904.—The Palaeochemistry of the Ocean in Relation to Animal and Vegetable Protoplasm. Trans. Canad. Inst., Vol. 7, pp. 535-562. — , 1910.—Inorganic Composition of the Blood. Proc. Roy. Soc. Lond. mmenine 2 oy 2 cocker y, of Body Fluids and Tissues. Physiol. Rev., Vol. 6,
pp. 816-357.
Marearia, R., 1931.—The Osmotic Changes in Some Marine Animals. Proc. Roy. Soc., 107 B, pp. 606-624.
XXxXil : PRESIDENTIAL ADDRESS.
MarsHALL, E. K., and SmitrH, H. W., 1930.—The Glomerular Development of the Vertebrate Kidney in Relation to Habitat. Biol. Bull., 59, p. 135.
Montl1, R., 1914.—La variabilita della pressione osmotica nelle diverse specie animali. Atti. Soe, ttal; Se. Nats. 53, De ool
NEEDHAM, J., 1931.—Chemical Embryology. Three Vols. Cambridge.
PANTIN, C. F. A., 1931.—Origin of the Composition of the body fluids in Animals. Biol. Rev. Camb., 6, No. 4, pp. 459-482.
Pearse, A. S., 1932.—Freezing Points of Blood of Certain Littoral and Hstuarine Animals. Pubn. No. 435, Carneg. Inst. Wash., pp. 93-102.
PorTiER, P., et Duvau, M., 1922.—Pression osmotique du sang de l’Anguille essuyée en fonction des modifications de salinité du milieu extérieur. C.R. Acad. Sci. Paris, U5, jo IolO,
QUINTON, R., 1897.—Hypothése de l’eau de mer, milieu vital des organismes élevés. C.R. Soc. Biol., Tome 49, pp. 935, 965 and 1063.
—————, 1904.—Communication osmotique chez le poisson Sélacien marin entre le milieu vital et le milieu extérieur. C.R. Acad. Sci., 139, pp. 995-7. See also Bull. Soc. Sci. Arcachon, 1904-5.
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RINGER, S., 1882.—Concerning the Influence exerted by each of the Constituents of the Blood on the Contraction of the Venitricle. Journ. Physiol., 3 and 4.
Ropirer, E., 1908.—Sur la Pression Osmotique du Sang et des Liquides Internes chez les Poissons Sélaciens. C.R. Acad. Sci., 131, p. 1008.
SCHLIEPER, C., 1929.—Ueber die Einwirkung niederer Salzkonzentrationer auf marine Organismen. JZ. vergl. Physiol., 9, pp. 478-514.
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ScHWABE, E., 1933.—Uber die Osmoregulation verschiedener Krebse (Malacostracen). Zeit. Vergl. Physiol., Band 19, Heft. 1, pp. 183-236.
SmMITH, H. W., 1931.—The Absorption and Excretion of Water. and Salts by the Elasmobranchs. I and II. Amer. Journ. Physiol., 98, No. 2, pp. 279-310.
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The Secretary (for Dr. G. A. Waterhouse, Honorary Treasurer) presented the balance-sheets for the year ended 28th February, 1935, duly signed by the Auditor, Mr. F. H. Rayment, F.C.A. (Aust.); and he moved that they be received and adopted, which was carried unanimously.
No nominations of other candidates having been received, the Chairman declared the following elections for the ensuing session to be duly made:
President: W. L. Waterhouse, M.C., D.Se.Agr., D.I.C. (Lond.).
Members of Council: E. C. Andrews, B.A., W. R. Browne, D.Se., E. Cheel,
A. G. Hamilton, Professor T. G. B. Osborn, D.Sc., and T. C. Roughley, B.Sc., HREZS:
Auditor: F. H. Rayment, F.C.A. (Aust.). A cordial vote of thanks to the retiring President was carried by acclamation.
XXX1li
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REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. III. By A. JEFFERIS TURNER, M.D., F.R.E.S. [Read 27th March, 1935.]
In my last instalment the description of 240, Borkhausenia xuthochroa, was accidentally omitted. I have failed to trace this specimen, consequently it must be omitted. I. have substituted another species for this number, and take the opportunity of describing several other species of that genus.
In the key to the genera is included a small group designated by Meyrick Scaeosophides, of which there are two genera known in Australia, together with a much larger group containing all the genera in which vein 7 of forewings terminates in the apex, with the exception of the Machimia group, which will be considered separately. Most of the genera are closely allied to Hulechria, and the distinctions are sometimes rather finely drawn. The criterion of the apical termination of vein 7 must not be applied too strictly. This vein seldom terminates in the mathematical apex, unless that is acute. When the apex is rounded, its termination is usually at the point where the upper end of the termen begins to curve; this we may term the anatomical apex. In some examples of several genera the termination may be just below this, and the determination of a species may then require the careful examination of several examples. In some instances the distinction may appear artificial, but in the present state of our knowledge we cannot dispense with it.
240. BoRKHAUSENIA BRACHYSTICHA, N. SP.
Bpaxvorcxos, with short streaks.
fg. 18-20 mm. Head and thorax grey-whitish sprinkled with fuscous. Palpi _ with second joint reaching base of antennae, terminal joint three-fifths, whitish with a few fuscous scales. Antennae grey with blackish rings, ciliations in male 1. Abdomen grey. Legs grey; posterior pair whitish. Forewings very narrow, costa slightly arched, apex pointed, termen very oblique; grey-whitish finely sprinkled with fuscous; stigmata represented by short longitudinal fuscous streaks, first discal at one-third, plical before it, second discal at two-thirds, dot- like, a streak above and between discals, and another between plical and second discal; cilia grey-whitish with a few fuscous points. Hindwings and cilia pale grey.
Tasmania: Lake St. Clair (2,000 ft.) in January; two specimens.
247. BoORKHAUSENIA LITHODES, N. SD. \LGwdns, Stone-coloured. . 6d. 17mm. Head whitish. Palpi with second joint reaching base of antennae, terminal joint three-fifths; fuscous. Antennae fuscous; ciliations in male 1. Thorax fuscous. Abdomen dark grey. Legs fuscous. Forewings with costa gently arched, apex pointed, termen very oblique; grey-whitish; markings and E
2, REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iil,
some irrcoration fuscous; a narrow basal fascia; a curved line from one-third costa to one-third dorsum, and another from two-thirds costa to tornus, stigmata included in these lines, which tend to be suffused and interrupted; cilia grey- whitish. Hindwings and cilia grey.
Tasmania: Hobart, in January; one specimen.
248. BoRKHAUSENIA TORNOSPILA, N. SD. TopvoomAos, With tornal spot.
9. 18 mm. Head and thorax whitish. Palpi reaching base of antennae, terminal joint three-fifths; fuscous, inner surface whitish. Antennae fuscous. Abdomen ochreous-grey; apices of segments and tuft whitish. Legs fuscous; posterior tibiae and rings on middle tibiae whitish. Forewings rather narrow, costa moderately arched, apex round-pointed, termen oblique; whitish with rather dense fuscous irroration and markings; first discal at one-fourth, plical before it, second discal about middle; a subterminal series of dots from beneath three-fourths costa around apex and termen to tornus; cilia grey, bases barred with fuscous. Hindwings and cilia whitish.
Tasmania: Mt. Wellington (2,500 ft.) in January; one specimen.
249. BorRKHAUSENIA BUTYREA, DN. Sp. Bouvrupeos, butter-coloured.
3, 9. 15-16 mm. Head yellow. Palpi with second joint exceeding base of antennae, terminal joint two-thirds; fuscous. Antennae fuscous, ciliations in male 13. Thorax fuscous; a posterior spot and tegulae except bases yellow. Abdomen fuscous; tuft ochreous. Legs fuscous; posterior pair ochreous. Fore- wings narrow, costa almost straight, apex rounded, termen obliquely rounded; ochreous-yellow; markings fuscous; first discal at one-third, plical beneath it, second discal at two-thirds, connected by a streak with tornus; a terminal fascia; terminal edge yellow; cilia yellow, on apex and tornus fuscous. Hindwings and cilia fuscous. Near B. cosmanthes; from this and its allies it may be distinguished by the absence of a basal fascia.
Western Australia: Kalamunda, near Perth, in December and January; five specimens received from Mr. W. B. Barnard, who has the type.
Key to Genera
i aindwines* with hyaline spatchi beneath Cell) Sryyaeren creel ciercteiaterc Neel etel oN nomena neal 2 Find wanes) swithouitehyaline spa t@liw eyeusweter-i-<ertcied nenletene) cliel-MelononslicPopcn- ficial Nai tena iaiei nema 3 2; PMOLE WANE Se with a 18s Oe Stale dO creceucueusiey clo peicier ciel oie ecient: chews 25. Neossiosynoecha MOKewingswathi 9 SEreey chute censveue tees sectors: eis me donsetel as aeee ceeea eI nae oeea ieee 26. Scaeosopha 3. Middle and posterior tibiae with whorls of projecting scales .................. 4 Middlevand posterior tibiae mnOt iSO) mieretne seine cicueiiel cnclemenceeken heen oi-ttel hel ite enema nene ms 6 Ai, INO UNS Woo, 7 Ehaycl (3 Gorineicleime seoasohoaavond0beauoucou00 se 27. Celeophracta Forewings, with. 7, -andi8) ‘stalked! soy0.58) ssn cysievs onic cheuete ensee oe Ge ROIS eee Ieee 5 D2- ANTENNAE KWIThOWe sDECCEIY Ay orecce ea neconys SUCKS me clin (olscelteeo hs eine ci eay scene a eeate 28. Trichomoeris Antennae with pecten. ei.c Aes eitetens. sistas eperahecelenerapsiiclaca tous couetterarensrencneeteee 29. Petalanthes 6. Tongue absent or rwa@inrentarye oie. Gan ciocs oe outs solve te) ove dopouorte: a elope ele ecnersitel MONG Re ER RnR Regie 1 Tongue normally Geveloped si eke scale ove lg cuekd eww www iat orm ae lane wilokav sl we ee RE eee 11 (. Hindwings with 3 and 4 well separate at origin .:....:2.....-l..-- 33. Gyrophylla Hindwinss® with 3--andl 4 (connate, Giwiecoe cha e 6 4 ee ecicinonm So ete EEE 8 So NGM bOI) Niall. 7 hovel } @oOMine@nGlOMme oacscdcboaocouscooddcuosobocuSN 30. Anomobela Poréewings with “7 ‘and’ 8 stalked sy Ve aac slneee aioer are at aie a nie tote ae na 9 9) Palpil iwith second’ joint tthickenedi ese ocelot eee nee ee eee 10 Palpimiwithssecond: sointyslender snare itte eT oeoreeee 31. Limothnes WHS) ASEVhol Vyielal, eeroecvioeMl soyime ON 5 cocccsbonocaconoudsosasoaosuoo‘e 32. Helactistis
Pelion Wrlioy Wesocbaeul aoe GEMS Jadecdccacodoudcsudueugdoonaosanoc 34. Phloeochroa
BY A. J. TURNER. 3
Jrinigrenere ome Eyal PAE Wawolrdrversl nisacloonoaonoonaocoDUodGHduoonOoOUnonoObOOS 12 AmMiSeore wionrae Arorel AAS SMO whi@iermeCl josacnodgnods00n00endboobooUeOanODO4HN GO 15 GNTGLEZIoiely ARAN CMON, MOMS Netgtoun eed O bl CO n ROR EROIIECEIC » Glofokarera Gite atcio camo oeLaraiure mec oton 13 TAO ASS AVL UEMCME Sen the cancer iereacnsuetaaet airs, 'o os istics cis: wile\/enel'e MMe abe ela detieray ou eitsr a) eatehionraite dalrolrewauetestouetrenrayl 14 ATAISAOAS WAGAOUE WACK, oncnonsomooonos5ogonoDO bo oMbomModb oo boon 35. Callimima AMLENNAIeH awilulapeCheNy ar. aederate alo lialacmene cls Ghsllcpe: opel encheieherevelocsterabeciarevclerstalter alls 36. Copriodes INORENMINESS WA WoLEES Che GOW coaaondconpcpodgnonbobunoboonoDDGOE 37. Piloprepes MOLES WAN SSSI Olle ees on firct ee epe tuciencbfane ie 2atesiicl:syaks) cs) AMP on (aylol apleeh cnelicneme denice olrciie 38. EHpipyrga Forewings with tufts of scales .............. Pini didkto olf Dane roto oeeraen 39. Trachypepla LOLS WATIES SIMO OUIMM Were eS et ene s nade tete rel ers «cls abe ielialta analrehecimep omretamelareiaiteaiah a. srvale (neha eve eet ale, tos stir 16 aD aVayars lose VAN Ob (CR CCEYS Be Sey tS ICR HCO ORCh Ik REE SOARS Paice Soins CLC LOMh OseaD TENOR ai amacd Die HenlD LAO Dro lola IY AM DNoAe ee N ANE GLOW HORSE. lo Gio oi CROeE EI ORO ORS ERMeaOT OI ICi EO aia-CI I cose o-cad Aro IONE Oo Ocha DID’ bio olo DG 20 PAtGnn alemmwlthOmltan CLOT aiieusyavcrusisuce sie cven fone.) aiteisey taustreticl Shoenrel otal em onsternes-ciiewan sttel chceseletiatts 42. Barea PNihenim ae miwaltlyn CCLETUm seer uence. cyte rend sey cea) isch g sevtedans susgtine peas eee ee re tate e reer chet ene reas toenetteioiene 18 Palpi with second joint dilated beneath, with rough scales towards apex ..........
ea EE ae ATTN cI Sree els Reh Ne eek earr eee ire talie elise ais snial Wivauleleeu al oun Mee duede MaRS eto alee teactione 40. Oenochroa Pap igen ote dil ated mt Ow ard Siva OX ire f feeli-papiais) elo al cian es eheistenhenomeleyeneirariicue ke temenemellel onesies 19 erlion Walla weal One ‘sileiCler Gossacogoeonunvcnodcodconsue0Gd 41. Placocosma IPeAllion weal Weremavloeyl Sfoubaye SWONE sootougoanodbonbouDpoGoonOcGDbOUOD 43. Hucryphaea OREN Vestn, 7 Eincl @Goneieleme Gro goocoedoocogsosboonDodbudoUo DOA SuOnaONOO 21. MORE walla SSenovile pa Greedy Che Sat SCALE Cues cie cers pcprevca ale slsscoroweuc pAUWel Cue Ra Rea cuenoncae ehe vols we eee Men cic errs ae 22 IMOLEWwINeSawalhe2wanadeor stale Gert, csi cicerel susicei cuavenscoreusenesienetencelente 51. Heliosteres MOnewiNsSiwilthiezman dma mse panacea ccicisicrelelscicicnsccnereicieie) ouch ceatne 55. Mermeristis Palpi with second joint in male dilated with loose hairs at apex ................ 23 TEE Waly o10NO Le LEKOYS: tyro kone ot Bo lg: Oso rOT NO AOC Oe ER ERED ERS RE Lis: Si IER CicLLrMeres bad ace chiara cure rato. Goriie cen 24 Palo Win weary! aoe Gemclere soonasoadaceoupoouucdoouDDOboOOUbOnS 44. Toptera Palpi with terminal joint moderately stout ...................... 45. Phriconyma PAMLCHIN VemhwAllLMOUt mu CHET! ware cle cic) ealiue solace oriole salve cama cortea a inliee tviciaiseiveltel Gnieclay egtapetratetcn CA aa arches 25 PANGENNE Ceawathnipe GEEMaa cu, sis tcrcien ces sanehe tS ais Gr se ek wera tare aaah Shy gees, Soe eg a acta an tae nn 30 MoOnewine smyth 2isand) sm stallke diy srs a scan «secre, sve ee steledeys Sa ein onehuecedstere 52. Actenotis MOGewAinsSsmwlthercwan davon OtustalKeds palais: -2-koncyeanieueker ere nets ea ero IC Cire 26 Palpi with second joint three times length of face ................. 46. Bathydoxa Ap iawlehseCOndey OIntnousexceedineutwicel face ae ieeeicieeeoe 27 MOLE WANES Withee onng eS ta LCE dine a teats Gul welayclacde sulci om Uateato Nee bey enn alta meses 47. Ancharcha LOGE WATE Sea vlt he Oper e Ce curser aa tes arepom sus syeliaicamusinen shisylepiei lass ia meite ar ieneu suis gaheoecrsueins Meronan Sizeenoushemsbematene 28 Palpi with second joint not reaching base of antennae ................ 49. Utidana PalpleawAthiasecondmjoint#neachinembase: of amtennalel manmec cl cieiececcreceiete oie 29 IPA on walla Recoil somone Gene soodoouvosnoovododoubdacdoobDU Sos 48. Locheutis PAyo WAIN SEooincl! sjyoybne Woaikeliewmecl GaccscaccusseucusodobonddobaouSoD 50. Allodapica HOnEWINe Simwithwo wands Stalked oasis cre ceye seas chelco tens cociereee seueieweroieiens 53. Hlaeonoma DOrRENVIMAES “Wikio 2 incl 8 imocessealaxol soso bacoococposdogccboddeG obo dGauccedoUS 31 Palpi with second joint extremely long, terminal joint less than half second ........
2 a'e, OS erg SiC OPEC A DORE! DIELORECC DY God ce ie Pan Urn E ALN ear Aime RIT meh iatt ste, 54. Hpithymema JEM OI F SOVOLGENKG)y Veet Gs, oer Ee ORTOrGY OCIA SOHC CES Sctltar Sa era pee RPRIRMEr rN An CRORA a Me ee Aber tal cea s rons Ewa as 32 IPAhon Wl ecology | sKorioe ewe, Ele WEIRD Ee EIS Goocasaucanunccuoosodscoauenbo 33 a pies wlLheteEminale| OlNntuslendersthroucshouity ser ciyoiiacicieciorcelcneioicichoneicieteneniemerane 35 LGTY WATE S ATCC OLE Mia trite earns) oe ha) lenjel ices rahe) See reat oe ake) Ce tone RO A TSO ANOS rata es 34 Elim eS melONSAte= OVALS matesie cane, etevche orien es cietia) Sualetei fo iduen evene dt iseeieteaeonon eines 58. Trachyntis EinGwin sss watherdtorar did) aStalked! es oc cisuctepeusy a: oneal steue) slcborel atoms seeeonueiene 56. Phloeocetes lahhnGhyioess sya ¢b eyavel by) SOREN) So odcoacnnacconcedboooggcd0DCS 57. Ischnophanes TEC AGHR ITER Y MEW ACEO ER C2) eh os BUA loneat ne ici ERO R MERC RSME RENEE ORE REM RET EMR eel OL AEa Gictnia Han Gente cc op loreearcea 36 Melo TI ChnN SSPRCLOM SATE =OMALC Ut receea lary aie ci scsuomero les loc la cere nc dle reuel allo evn lore EAR RET ne eas act cl AREAS 38 Hindwings with 5 connate or stalked with 4 .................. 59. Hlaphromorpha TB OEANON TIO Mapiatidale Chin celravol Gyr yey ofe meee) ighy eee Enis BME cree no Ss oes OO B bin dann oa eas 37 indwines! with! 5) strongly, approximated to 6 ...5..5.+5.4.sodso+ eee 60. Asthenica indwines) withe 5: not approximated to 6 22s... 0ss4ss0esee se se 61. Macronemata Palpi with second joint not reaching base of antennae ........... 62. Brachyzancla PaAlpluamithasecondmyolmntncachinenbpase On antennaes smc seeecic ed ssiciieceeiena one 39 Palpi with long rough hairs on posterior surface of second joint .. 63. Anomozancla IPA Non, seeAKEL MONG: SIG VoWNT a MeI Nase cee ole cy ola ee ean Re PREM Sette A err. Sono cd Gilead anckols euemeie vais iors 40 Palpi with second joint more than twice length of face ........ 64. Phanerozancla
Palpi with second joint less than twice length of face .............. 65. Hulechria
4 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iii,
25. Gen. NrossiosyNoEcHA Turn.
Trans. Ent. Soc., 1923, p. 171. Type, T. scatophaga.
Tongue present. Palpi ascending, recurved, smooth-scaled; second joint moderately long; terminal joint much shorter than second. Antennae with basal pecten; in male moderately ciliated. Forewings with 2 and 3 stalked, 5 absent, 7, 8, 9 stalked, 7 to termen. Hindwings with 3 and 4 stalked, 5 from middle of cell; a triangular basal hyaline area beneath cell.
This and the following genus belong to Meyrick’s Scaeosophides characterized by the hyaline patches on the hindwings.
' Two species: 250, scatophaga Turn., Tr. EH. S., 1928, p. 172 (Coen, N.Q.).— 251, agnosta, n. sp.
251. NrEOSSIOSYNOECHA AGNOSTA, N. SD. ayvworos, unknown.
6. 35 mm. Head fuscous sprinkled with whitish. Palpi with second joint reaching base of antennae, terminal joint two-fifths; fuscous sprinkled with whitish. Antennae fuscous; ciliations in male 2. Thorax dark fuscous sprinkled with whitish. Abdomen ochreous-fuscous. Legs fuscous. with whitish rings; pesterior pair grey-whitish. Forewings narrow, suboval, costa slightly arched, apex rounded, termen obliquely rounded; dark fuscous with fine whitish irroration appearing grey; veins more whitish; stigmata blackish, first discal at one-third, plical beneath it, both these are narrow and streak-like, second discal before two-thirds; cilia fuscous sprinkled with whitish. Hindwings elongate; whitish-grey; cilia whitish-grey.
Structurally exactly as the type species, but with narrower wings. We should much like to know the larval habits of this species.
Queensland: Toowoomba, in October; one specimen received from Mr. W. B. Barnard, who has the type.
26. Gen. ScarosopHa Meyr.
Exot. Micro., i, p. 254. Type, 8S. percnaula Meyr., from India.
Tongue present. Palpi with second joint not reaching base of antennae; terminal joint shorter than second. Antennae with basal pecten; in male shortly and unevenly ciliated. Forewings with slight tufts of scales; 7 and 8 stalked, 7 to apex, 9 free. Hindwings elongate-ovate; cell less than one-half; a triangular basal hyaline patch beneath cell; 5 from below middle.
There are two Indian species. Lower’s name for the Australian species has a few months’ priority. In one of my examples 7 and 8 are coincident in one forewing.
252, epileuca Low., Tr. R.S. S. Aust., 1901, p. 94 (= mitescens Luc., P.R.S.Q., USO, ay Bile
27. Gen. CELEOPHRACTA, 0.2. Kn\eoppaxtos, fiery-edged.
Palpi moderately long, recurved, ascending; second joint reaching base of antennae, slightly thickened, with smoothly appressed scales; terminal joint shorter than second, slender, acute. Antennae without pecten; in male with rather long ciliations. Middle and posterior tibiae with median whorls of projecting hairs. Forewings with 7 and § coincident. Hindwings with 3 and 4 coincident.
This and the two following genera form a peculiar little group. Type, C. corusca.
BY A. J. TURNER. 5
253. CELEOPHRACTA CORUSCA, Nl. Sp. coruscus, shining.
3, 9. 15-16 mm. Head fuscous; posterior margin ochreous-yellow. Palpi orange-ochreous; terminal joint’ three-fifths; fuscous. Antennae blackish; ciliations in male 2. Thorax dark fuscous; tegulae and a posterior spot shining brassy. Abdomen fuscous; apical segments fuscous-ochreous with whitish apices. Legs fuscous; tibiae and tarsi with whitish rings. Forewings narrow, costa nearly straight, apex rounded, termen oblique; blackish uniformly irrorated with slender whitish scales; some basal ochreous irroration; ochreous dots in disc at two-thirds and on fold; terminal edge and cilia brassy with metallic reflections; cilia on apex and tornus grey. Hindwings and cilia grey.
South Australia: Adelaide (Coll. Lower); Mt. Lofty (Blackwood; J. D. O. Wilson) in October; four specimens.
254. CELEOPHRACTA HYPEREPHANA, DN. SD. Urepydavos, CONSpicuous.
6. 15-16 mm. Head fuscous; posterior margin orange-ochreous. Palpi orange- ochreous; terminal joint four-fifths, fuscous. Antennae fuscous; ciliations in male 2. Thorax fuscous with brassy lustre. Abdomen fuscous with a broad post-median ochreous band; underside mostly ochreous-whitish. Legs fuscous; tibiae and tarsi with ochreous-whitish rings. Forewings narrow, costa nearly straight, apex obtuse, termen oblique; brown-fuscous uniformly irrorated with narrow ochreous-whitish scales; a pale ochreous dorsal spot immediately followed by a tornal fuscous spot; terminal edge and cilia brassy with metallic reflections; cilia on apex and tornus grey. Hindwings orange-ochreous; terminal half fuscous; cilia fuscous.
Western Australia: Mundaring near Perth, and Busselton, in October; two specimens received from Mr. G. M. Goldfinch, who has the type.
28. Gen. TricHomorERIs Meyr.
Hxot. Micro., i, p. 156. Type, T. amphichrysa.
Tongue present. Palpi with second joint reaching base of antennae, terminal joint shorter than second. Antennae without basal pecten. Middle and posterior tibiae with median whorls of rough hairs. Forewings with 7 and 8 stalked, 7 to apex. Hindwings elongate-ovate; neuration normal.
Two species: 255, amphichrysa Meyr., Exot. Micro., i, p. 156 (Darwin; Cairns) .—256, heterochrysa Meyr., Arkiv. f. Zool., xiv (15), 6 (Atherton).
29. Gen. PETALANTHES Meyr.
Proc. Linn. Soc. N.S.W., 1883, p. 335. Type, P. sphaerophora Meyvr.
Tongue present. Palpi with second joint reaching base of antennae, terminal joint rather shorter than second. Antennae with basal pecten of few but long scales; in male with tufts of long cilia. Middle and posterior tibiae with median whorls of rough projecting hairs. Forewings with tufts of scales; 7 and 8 stalked, 7 to apex. Hindwings elongate-ovate; neuration normal.
Some of the species have remarkably spotted hindwings.
Five species: 257, sphaerophora Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 335 (Toowoomba, Sydney, Katoomba).—7258, diploxantha Meyr., Exot. Micro., i, p. 236 (Newcastle).—259, hexastera Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 336 (Brisbane to Melbourne) .—260, microphrica, n. sp. (Mittagong).—261, periclyta Meyr., ibid., p. 337 (Brisbane, Toowoomba, Sydney).
6 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iii,
260. PETALANTHES MICROPHRICA, 0. SD.
futkpopptxos, minutely rippled.
6g. 14 mm. Head and thorax fuscous. Palpi white laterally, fuscous anteriorly and posteriorly. Antennae blackish with fine whitish annulations; ciliations in male 4. Abdomen and legs fuscous. Forewings dilated, costa straight, apex rounded, termen rounded, slightly oblique; whitish-ochreous evenly traversed by five sinuate fuscous transverse strigulae; a whitish-ochreous subcostal dot near base; a narrow transverse dark fuscous fascia at one-fourth, imme- diately followed by a median whitish-ochreous dot; a transverse median whitish bar, edged with dark fuscous and with a dark fuscous dot on lower end; a second dark fuscous fascia at three-fourths, broader on costa; cilia whitish-ochreous with two fuscous lines. Hindwings similar, but fuscous fasciae broadly suffused, separated by a moderate whitish fascia; cilia grey with a sub-basal line.
New South Wales: Mittagong, in November; one specimen in Coil. Goldfinch.
30. Gen. ANOMOBELA, N.g. avowoBedos, with unusual palpi.
Tongue absent. Palpi moderately long, ascending, recurved; second joint reaching base of antennae, thickened with appressed scales, rough towards apex anteriorly; terminal joint one-fourth or less, slender, acute. Antennae with basal pecten; ciliations in male moderately long. Thorax smooth. Forewings with 7 and 8 coincident. Hindwings elongate-ovate, neuration normal.
Allied to the following genus; the affinities of the two genera are uncertain.
262. ANOMOBELA PLICILINEA, N. SD. plicilineus, with a line on fold.
6. 15 mm. 92. 18 mm. Head whitish. Palpi with terminal joint in male one-fourth, in female one-sixth; fuscous, most of inner surface and apex of second joint whitish. Antennae grey-whitish; ciliations in male 1%. Thorax fuscous; tegulae grey-whitish. Abdomen grey-whitish with some ochreous suffusion. Legs fuscous with ochreous rings; posterior pair ochreous-whitish. Forewings narrow, oval, costa rather strongly arched, apex pointed, termen extremely oblique; whitish with three suffused pale fuscous fasciae; first broad, basal; second broad on costa from one-third to two-thirds, becoming narrower towards dorsum before middle, ill-defined posteriorly; third narrow, from three-fourths costa to before tornus, acutely angled outwards in middle; stigmata shortly linear, blackish, first discal shortly before middle, plical before it, second discal at two-thirds trans- verse; cilia whitish. Hindwings and cilia whitish-grey.
North Queensland: Cape York, in June; two specimens received from Mr. W. B. Barnard, who has the type.
31. Gen. LIMOTHNES, D.g. Aywobvyns, Starving.
Tongue absent. Palpi smooth, slender, ascending, recurved; second joint not reaching base of antennae; terminal joint shorter than second. Antennae with basal pecten; in male moderately ciliated. Abdomen stout. Forewings with 7 to apex. Hindwings ovate-lanceolate, 5 from below middle.
263. LIMOTHNES LEUCOTOMA, Nl. SDP. AeuKoTomos, divided by white. dg. 18-19 mm. Head fuscous; face whitish. Palpi with terminal joint three- fifths; fuscous, apex of second joint and base and apex of terminal joint whitish.
BY A. J. TURNER. 7
Antennae whitish; ciliations in male 13. Thorax whitish anteriorly and posteriorly suffused with fuscous. Abdomen ochreous-brown; apices of segments and tuft pale grey. Legs whitish-ochreous; anterior and middle pairs with some fuscous suffusion. Forewings rather narrow, oval, costa moderately arched, apex pointed, termen extremely oblique; fuscous; a rather narrow white fascia from one-third costa to one-third dorsum; terminal area suffused with whitish; a discal dot in middle, and two dots often confluent at two-thirds, fuscous; cilia whitish. Hind- wings and cilia pale grey.
North Queensland: Cape York, in October and December; six specimens received from Mr. W. B. Barnard, who has the type.
32. Gen. HcnActistis Meyr.
Exot. Micro., i, p. 134. Type, EH. byrseuta Meyr.
Tongue weakly developed and rudimentary. Palpi long, ascending, recurved; second joint reaching base of antennae, much thickened with appressed scales, slightly rough anteriorly; terminal joint much thickened like second joint, apex obtusely pointed. Face with a pair of strong tufts curving over eyes. Antennae without basal pecten; ciliations in male moderately long. Thorax smooth. Fore- wings with 7 to apex or nearly so. Hindwings elongate-ovate; neuration normal.
Also represented in New Guinea by the type species, which has additional male secondary characters in the posterior tibiae and tarsi.
264. HcCLACTISTIS ANISOPASTA, 0. SD. avicomaortos, unevenly sprinkled.
6. 17-20 mm. Head ochreous-whitish. Palpi ochreous-whitish with a few fuscous scales. Antennae ochreous-whitish; ciliations in male 134. Thorax ochreous-whitish. Abdomen and legs whitish-ochreous. Forewings not dilated, costa strongly arched, apex round-pointed, termen very obliquely rounded; ochreous-whitish with patchy fuscous irroration; stigmata dark fuscous, first discal at one-third, plical beyond it, second discal at two-thirds; suffused fuscous patches near base, above mid-dorsum, and on costa at three-fifths and four-fifths, from the last a more or less curved line to tornus; cilia ochreous-whitish with some fuscous points. Hindwings grey; cilia grey-whitish.
North Queensland: Cape York, in October and November; six specimens received from Mr. W. B. Barnard, who has the type.
33. Gen. GYROPHYLLA, N.g. yupopdvAdos, with rounded wings.
Tongue absent. Palpi short, slender, not reaching middle of face; second joint with loose hairs beneath; terminal joint shorter than second, rather stout. Antennae without basal pecten; ciliations in male short. Thorax with a small posterior crest. Forewings with 2 from near angle, 7 and 8 coincident to apex. Hindwings with 3, 4, 5, 6, 7 nearly equidistant, parailel.
265. GYROPHYLLA EUMETRA, N. Sp. evmeTpos, well measured.
6. 22mm. Head whitish-ochreous. Palpi fuscous. Antennae grey; ciliations in male two-thirds. Thorax grey-whitish. Abdomen grey; tuft grey-whitish. Legs fuscous; posterior pair whitish. Forewings oval, costa strongly arched, apex rounded, termen obliquely rounded; grey-whitish; costal and terminal edge grey; cilia pale grey. Hindwings grey; cilia grey-whitish.
8 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iii,
New South Wales: Sydney (Manly) in March (G. H. Wyld); one specimen. Type in Coll. Goldfinch.
34. Gen. PHLOEOCHROA, N.g. @Aovoxpoos, coloured like bark.
Tongue absent. Palpi with second joint exceeding base of antennae, thickened throughout with appressed scales, dilated and slightly rough at apex; terminal joint less than one-half, slender, acute. Antennae with basal pecten. Forewings with 2 and 3 connate, 7 to apex. Hindwings elongate-ovate; neuration normal.
266. PHLOEOCHROA POLYRRHABDA, Dl. SD.
modveoaeoc, many-streaked.
2. 84 mm. Head and thorax whitish. Palpi with terminal joint one-third; grey, internal surface whitish. Antennae grey. Abdomen grey; tuft grey-whitish. Legs grey; posterior pair grey-whitish. Forewings elongate, costa slightly arched, apex pointed, termen very oblique; grey; costa towards base and dorsum broadly suffused with brown-whitish; disc sprinkled with a few fuscous scales; fine fuscous interneural streaks in post-median area; cilia grey. Hindwings and cilia pale grey.
New South Wales: Sydney (Manly) in March (G. H. Wyld); type in Coll. Goldfinch.
35. Gen. CALLIMIMA, N.g. KaAAcutwos, a beautiful mimic.
Tongue present. Palpi with second joint reaching or exceeding base of antennae, thickened with appressed scales, expanded and sometimes forming a triangular tuft at apex; terminal joint shorter than second, slender or rather stout, acute. Antennae without basal pecten; ciliations in male moderately long. Thorax smooth. Anterior tibiae and tarsi strongly dilated. Forewings with costal tuft; 7 to apex. Hindwings ovate; 5 curved from below middle of cell.
Type, OC. lophoptera. Near Copriodes, differing in the palpi and absence of antennal pecten.
Two Species: 267, lophoptera Low., Tr.R.S.S.Aust., 1894, p. 96 (Brisbane to Allyn River, N.S.W.).—268, daedalma, n. sp. (Cairns).
268. CALLIMIMA DAEDALMA, D. SD. dacdadua, a work of art.
2. 20 mm. Head whitish. Palpi whitish, outer surface of second joint fuscous towards base. Antennae grey, towards base whitish. Thorax grey- whitish. Abdomen ochreous-whitish. Forewings with costa strongly arched, bearing a strong median tuft, apex rounded, termen obliquely rounded; dark fuscous; costal edge except at base pale rosy; a well-defined whitish line along costa, continued around apex and termen to tornus; a broad ridge of whitish scales, anteriorly brown mixed with fuscous, from base to one-fourth dorsum; a curved whitish line from midcosta to before tornus, sharply defined and slightly waved on anterior edge; a narrow fuscous terminal line thickened on veins; cilia grey-whitish, on costa pale rosy. Hindwings grey; terminal edge whitish; cilia pale grey.
North Queensland: Kuranda, in May; one specimen.
36. Gen. CopriopEs Turn. Proc. Linn. Soc. N.S.W., 1916, p. 339. Type, OC. aristocratica.
BY A. J. TURNER. 9
Tongue present. Palpi with second joint reaching base of antennae, slender and smooth, or slightly thickened and rough anteriorly beyond middle; terminal joint shorter than second, slender, acute. Antennae with strong basal pecten; ciliations in male moderate or long. Thorax smooth. Anterior tibiae and tarsi strongly dilated. Forewings with costal tuft or with tufts of raised scales in disc; 7 to apex. Hindwings ovate or elongate-ovate; neuration normal.
In C. aristocratica the forewing has a peculiar shape (a specific adaptation for mimetic purposes) so that 7 runs apparently to costa. In C. anassa the costal tuft is scarcely developed, in ©. gelidella it appears to be absent, but is replaced by raised scales in disc. The genus differs from Piloprepes in the absence of a thoracic crest.
Hight Species: 269, aristocratica Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 1598 (Yeppoon to Fernshaw).—270, perinephela, n. sp. (Lismore).—271, hypsilopha, n. sp. (Cunnamulla, Dalby).—272, polynephela, n. sp. (Dimboola, Vic.).—273, anguicula Meyr., Exot. Micro., i, p. 133 (Brentwood, Vic.).—274, anassa Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 1597 (Toowoomba to Melbourne) .—275, glaucaspis Turn., T7r.R.S.S.Aust., 1896, p. 19 (Brisbane) —276, gelidella Wlk., xxix, p. 766. = lucasii Turn., ibid., 1896, p. 19 (Darwin, Brisbane, Sydney).
270. CoOPRIODES PERINEPHELA, N. Sp.
mepivepedos, Clouded.
®. 18 mm. Head white. Palpi white; apex of terminal joint fuscous. Antennae grey. Thorax fuscous; tegulae white. (Abdomen missing.) Legs white; tarsi with blackish rings. Forewings with costa bisinuate, bearing a tuft before middle, apex round-pointed, termen obliquely rounded; 7 to apex; white; blackish dots on base of costa and dorsum, and paired dots in disc just beyond base; a fine fuscous line from one-fourth costa, outwardly oblique, angled, and continued along fold for a short distance; again angled and curved outwards to mid-dorsum; a cloudy incomplete fuscous fascia, inwardly curved from beneath three-fourths costa to tornus, strongly excavated anteriorly, less so posteriorly; a similar marginal fascia from three-fourths costa to midtermen, but not touching termen; an interrupted dark fuscous terminal line; cilia whitish, barred with fuscous around apex. Hindwings grey; cilia grey, on dorsum whitish.
Nearest C. aristocratica.
New South Wales: Rous, near Lismore, in November; one specimen received from Mr. V. J. Robinson.
271. COoPRIODES HYPSILOPHA, 0. SD.
bYirogos, high-crested.
do. 1420 mm. Head whitish; apices of side-tufts fuscous-brown. Palpi whitish. Antennae grey, towards base whitish; ciliations in male 14. Thorax ochreous-whitish; anterior edge and two posterior dots dark fuscous. Abdomen fuscous-brown; dorsum of first two segments whitish. Legs whitish; middle tibiae fuscous with whitish rings. Forewings suboval, costa strongly arched, witi a slight indication of a tuft beyond middle, apex rounded, termen very obliquely rounded; ochreous-whitish; a raised tuft of scales in middle at one-third, and another at mid-dorsum, both dark fuscous at apices; some dark fuscous irroration in median area; two raised tufts arranged obliquely in disc beyond middle, dark fuscous; a fine crenulate fuscous line from costa beyond middle to tornus; beyond this line clear white except a small very faint subapical cloud; cilia
10 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iii,
white. Hindwings whitish-ochreous, suffused with grey beyond middle; cilia whitish-ochreous, around apex grey. Queensland: Cunnamulla; Jandowae near Dalby; two specimens.
272. COPRIODES POLYNEPHELA, 0. SDP. mo\vvepedos, Much clouded.
do. 16 mm. Head whitish. (Palpi missing. Antennae imperfect.) Thorax whitish, central area and a posterior spot fuscous. Abdomen pale ochreous. Legs whitish. Forewings suboval, costa strongly arched, apex rounded, termen obliquely rounded; whitish; pale fuscous basal markings, namely, a narrow curved transverse line near base, a dot on costa at one-sixth, a large spot on dorsum at one-fourth, connected with fascia on dorsum, limited by fold, and a smaller spot in disc above fold; an ochreous-grey-whitish fascia from two-fifths costa to mid- dorsum, dilated in middle, constricted on fold, dilated again on dorsum; an irregular fascia from mid-costa to tornus, at first pale fuscous, connected with median above middle, becoming ochreous-grey-whitish in disc, but pale fuscous towards tornus, sharply indented posteriorly above tornus, posteriorly edged with black except towards margins, strongly outwardly curved, but indented above middle; a subapical bluish-fuscous crescent; cilia whitish with an apical fuscous dot, on tornus grey. Hindwings grey; cilia grey, towards apex whitish.
Victoria: Kiata, near Dimboola, in November; one specimen. Type in Coll. Lyell.
37. Gen. Prnopreres Meyr.
Proc. Linn. Soc. N.S.W., 1883, p. 365. Type, P. aemulella.
Tongue present. Palpi with second joint reaching base of antennae, slightly thickened with appressed scales; terminal joint shorter than second, slender, acute. Antennae with basal pecten; in male with moderate ciliations. Thorax with a strong posterior crest. Anterior tibiae and tarsi slightly dilated. Fore- wings with tufts of scales; 7 to apex. Hindwings elongate-ovate; neuration normal.
Readily distinguished from Copriodes by the strong thoracic crest. With the two preceding genera it forms a natural group.
Two Species: 277, aemulella Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 366 (Darwin, Cape York to Melbourne) .—278, antidoxa Meyr., ibid., 1888, p. 1599 (West Victoria, Adelaide. W.A.: Cunderdin, Beverly). ;
38. Gen. Hprpyrca Meyr.
Proc. Linn. Soc. N.S.W., 1884, p. 791. Type, #H. agaclita.
Tongue present. Palpi with second joint reaching base of antennae, thickened with rough scales anteriorly; terminal joint shorter than second, slender, acute. Antennae without basal pecten; in male moderately ciliated. Thorax not crested. Anterior tibiae and tarsi slightly dilated. Forewings smooth; 7 to apex. MHind- wings elongate-ovate; neuration normal.
EH. hemiphanes might be an aberration of H. agaclita, but I do not think so. It differs in the wholly fuscous thorax, basal fascia on forewings, darker apical half of wing without yellowish spot, and purple-fuscous cilia.
279, agaclita Meyr., Proc. Linn. Soc. N.S.W., 1884, p. 791 (Cairns to Brisbane and Milmerran) —280, hemiphanes Turn., Tr. R.S.S. Aust., 1917, p. 58 (Brisbane).
39. Gen. TRACHYPEPLA Meyr.
Proc. Linn. Soc. N.S.W., 1883, p. 367. Type, 7. euryleucota Meyr. from New Zealand.
BY A. J. TURNER. 11
Tongue present. Palpi with second joint not reaching, reaching, or exceeding base of antennae, thickened with appressed scales; terminal joint shorter than second, slender, acute. Antennae with basal pecten; ciliations in male moderate or long. Thorax sometimes with a small posterior crest. Forewings with tufts of raised scales: 2 usually separate, rarely connate or even stalked with 3, 7 to apex. Hindwings elongate-ovate or broadly lanceolate; neuration normal.
This genus is confined to Australia and New Zealand. Meyrick records twenty species from the latter region. It presents considerable variation in structure. The Australian species might be divided into two groups: (1) those with lanceolate hindwings and second joint of palpi not reaching antennae, (2) those of larger size with elongate-ovate hindwings and second joint of palpi exceeding base of antennae, but the New Zealand species are intermediate. It appears inadvisable to break up a natural genus of moderate size into artificial genera, which grade into each other.
Fifteen Species: 281, atrispersa Turn., Proc. Linn. Soc. N.S.W., 1916, p. 347 (Brisbane to Sydney).—j7282, charierga Meyr., ibid., 1888, p. 1566 (Bathurst; Deloraine, Tas.; Perth, W.A.).—283, phaeolopha, n. sp. (Nambour, Brisbane, Toowoomba, Stanthorpe).—284, poliochroa Turn., Tr.R.S.S.Aust., 1898, p. 208 (Brisbane, Mt. Tambourine, Toowoomba, Stanthorpe).—285, lasiocephala Low., ibid., 1916, p. 540 (Dalby).—286, stenota Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 567 (Sydney; Perth, W.A.).—7287, hemicarpa Meyr., ibid., 1887, p. 954 (North Tasmania).—288, melanoptila Meyr., ibid., 1883, p. 370 (Brisbane, Sydney) .— 289, diplospila, n. sp. (Toowoomba) .—290, picimacula, n. sp. (Sydney).—291, capsellata Meyr., Exot. Micro., i, p. 157 (Beaconsfield, Vic.; Tasmania, Mt. Lofty) .— 292, glebifera Turn., P.R.S. Tas., 1926, p. 142 (Tasmania) —293, dasylopha Low., Tr. R.S.S.Aust., 1920, p. 61 (Dalby).—294, haemalea Turn., Proc. Linn. Soc. N.S.W., 1916, p. 347. = plinthinopa Meyr., Exot. Micro., ii, p. 368 (Hidsvold, Brisbane, Toowoomba) .—295, peplasmena, n. sp. (Sydney).
283. TRACHYPEPLA PHAEKOLOPHA, TD. Sp. patodogos, dark crested.
3, °. 10-15 mm. Head white. Palpi whitish; second joint with basal half and a subapical ring fuscous. Antennae grey-whitish; in male slightly serrate, ciliations one-half. Thorax white. Abdomen pale grey; tuft ochreous-whitish. Legs fuscous mixed with whitish; posterior pair ochreous-whitish. Forewings narrow, costa gently arched, apex rounded; white with some pale ochreous-grey irroration; dark fuscous costal dots at one-fourth and beyond middle; a large tuft of raised scales on mid-dorsum extending half across wing, dark fuscous; cilia whitish finely sprinkled with fuscous. Hindwings lanceolate; grey-whitish; cilia grey-whitish.
Queensland: Eumundi, in November; Brisbane, in August; Toowoomba, in September and October; Stanthorpe; six specimens.
289. TRACHYPEPLA DIPLOSPILA, N. SD. Ourd\oortAos, two-spotted.
6, ©. 19-20 mm. Head and thorax white. Palpi with second joint much exceeding base of antennae, terminal joint three-fifths; white, base of second joint dark fuscous externally. Antennae whitish; ciliations in male two-thirds. Abdomen whitish-ochreous. Legs whitish. Forewings rather narrow, suboblong, costa gently arched, apex rounded, termen very obliquely rounded; white; some pale grey suffusion towards apex and on dorsum; two dark fuscous spots with
12 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iii,
large scales, being first discal at one-fourth and plical well beyond it, elongate; second discal sometimes indicated by a minute dot beyond middle; cilia whitish sprinkled with grey. Hindwings and cilia grey.
Queensland: Toowoomba, in November; two specimens received from Mr. W. B. Barnard, who has the type.
290. TRACHYPEPLA PICIMACULA, N. Sp.
picimaculus, blotched with pitch-black.
6. 22-24 mm. Head and thorax fuscous-brown. Palpi pale brown, on outer surface sprinkled with blackish; terminal joint three-fourths. Antennae fuscous; ciliations in male 1. Abdomen grey. Legs fuscous with whitish rings; posterior pair mostly whitish. Forewings slightly dilated, costa rather strongly arched, apex round-pointed, termen obliquely rounded; fuscous-brown with blackish markings; a strong line on fold from near base to two-thirds; a small irregular blotch, sometimes reduced to a short line, beneath costa at one-third; a second small blotch above middle emitting a posterior Y-forked line, its extremities approaching costa and dorsum, sometimes obscured by blackish irroration; some- times two subterminal dots below its lower extremity; a series of terminal dots; cilia pale fuscous sprinkled with blackish. Hindwings with 5 from middle; pale grey; cilia grey-whitish.
New South Wales: National Park near Sydney, in August; two specimens received from Mr. G. M. Goldfinch, who has the type.
295. TRACHYPEPLA PEPLASMENA, N. SDP. memAauevos, dissembling.
6. 22 mm. Head brownish. Palpi with second joint exceeding base of antennae, terminal joint three-fifths; whitish, second joint with irroration and a subapical ring fuscous; terminal joint fuscous except apex. Antennae fuscous; ciliations in male 1. Thorax fuscous. Abdomen grey; tuft ochreous-whitish. Legs fuscous with ochreous-whitish rings; posterior pair mostly ochreous-whitish. Forewings strongly dilated, costa strongly arched, apex rounded, termen obliquely rounded; whitish with fine fuscous irroration, appearing grey; markings dark fuscous; a subdorsal ridge of raised scales from base to one-fifth; a tuft of whitish scales above two-fifths dorsum; first discal at one-fourth, second at middle, trans- versely elongate; plical represented by a short line on fold, partly overlapped by subdorsal tuft; a stout inwardly oblique line from costa before apex, very acutely angled above middle of dise, continued by a slender line to dorsum before tornus, connected by an interrupted longitudinal streak from angle with second discal; a series of dots on termen and apical fourth of costa; cilia fuscous, apices whitish. Hindwings and cilia grey-whitish.
New South Wales: Heathcote near Sydney, in August; one specimen received from Dr. R. J. Tillyard.
40. Gen. OfNocHROA Meyr.
Proc. Linn. Soc. N.S.W., 1883, p. 327. Type, O. laetella.
Tongue present. Palpi with second joint reaching or not reaching base of antennae, thickened with appressed scales, and rough projecting scales towards apex anteriorly; terminal joint shorter than second, slender, acute. Antennae with basal pecten; in male shortly or moderately ciliated. Thorax with a small posterior crest. Forewings with 2 from well before angle, 7 to apex. Hindwings elongate-ovate; 5 from below middle.
BY A. J. TURNER. 13
Fifteen Species: 296, thermistis Low., Tr. R.S.S. Aust., 1896, p. 166 (Victoria, Tasmania). = atradelpha Low., ibid., 1908, p. 221.—297, laetella (lactella) WIk., xxix, p. 648 (Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 328) (Brisbane to Melbourne).—298, ochrosoma Turn., Tr.R.S.S.Aust., 1896, p. 13 (Brisbane, Toowoomba) .—299, lepida, n. sp. (Roma).—300, zophocosma, n. sp. (Roma).— 301, molybdoptera, n. sp. (Charleville). —302, suffulva, n. sp. (Atherton, Banana, Q., Birchip).—303, gnophodes Turn., ibid., 1896, p. 14 (Brisbane, Melbourne) .— 304, dinosema Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 1575 (Victoria, Adelaide) .— 305, zalotypa, n. sp. (W.A.: Denmark).—306, endochlora Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 329 (Warragul, Quorn, Wirrabara, Mt. Lofty) —307, dystena, n. sp. (Brisbane) .—308, homora Meyr., Tr. R.S.S. Aust., 1902, p. 152 (Sydney, Hobart) .— +309, heptarcha Meyr., Proc. Linn. Soc. N.S.W., 1888, p. 1576 (W.A.: Geraldton) .— 310, iobaphes Meyr., ibid., 1883, p. 330 (Brisbane to Tasmania, Mt. Lofty).
299. OENOCHROA LEPIDA, N. SD.
lepidus, pleasing.
°. 20-22 mm. Head and thorax blackish irrorated with grey-whitish. Palpi with second joint reaching base of antennae, terminal joint three-fifths; grey- whitish irrorated with blackish. Antennae blackish. Abdomen grey; apices of segments ochreous-whitish. Legs blackish; rings on tibiae and tarsi, and hairs on posterior tibiae ochreous-whitish. Forewings oval, costa strongly arched, apex pointed, termen very obliquely rounded; grey-whitish with some blackish irrora- tion and markings; a rather large triangular sub-basal costal spot; an outwardly curved line from one-fourth costa to one-fourth dorsum, giving off in middle a fine or interrupted line to tornus; a thicker line from two-thirds costa to tornus, expanded in middle to contain a grey-whitish spot; a series of dots or short streaks on apical one-third of costa and tornus; cilia grey. Hindwings ochreous- yellow; apex suffused with grey; cilia grey.
Queensland: Roma, bred from larvae on leaves of a broad-leaved Hucalyptus, two specimens emerging in Brisbane in August.
300. OFNOCHROA ZOPHOCOSMA, Nl. SD. Somoxoouos, darkly adorned.
& 9. 16-19 mm. Head grey-whitish sprinkled with dark fuscous. Palpi with second joint slightly expanded with rough scales towards apex anteriorly; terminal joint four-fifths; dark fuscous sprinkled with grey-whitish. Antennae dark fuscous; ciliations in male 1. Thorax with small posterior crest; dark fuscous sprinkled with grey-whitish. Abdomen grey; tuft ochreous-whitish. Legs fuscous with grey-whitish rings; hairs on posterior tibiae ochreous-whitish. Forewings narrow, oval, costa gently arched, apex round-pointed, termen very oblique; grey- whitish; markings and some irroration dark fuscous; a rather iarge suffused basal patch; a broad dorsal streak separated, sometimes incompletely, from basal patch at one-fourth to tornus, containing some grey-whitish scales before tornus; a discal dot at two-thirds; a costal subapical spot sometimes connected by a fine line with tornus; a series of ill-defined spots or short streaks on apical fourth of costa and termen; cilia grey, bases and extreme apices partly whitish. Hind- wings and cilia grey.
Queensland: Roma, feeding on the same Hucalyptus leaves as the preceding species, emerging in Brisbane in July and August; six specimens.
,14 REVISION OF AUSTRALIAN LEPIDOPTERA. OECOPHORIDAE. iii,
301. OENOCHROA MOLYBDOPTERA, MN. SD. podvBdortepos, leaden-winged.
9. 18 mm. Head dark fuscous; side tufts whitish. Palpi with second joint exceeding base of antennae, thickened with rough scales towards apex anteriorly, terminal joint three-fourths; fuscous. Antennae fuscous. Thorax dark fuscous. Abdomen whitish-grey. Legs fuscous; posterior pair whitish. Forewings elongate-oval, costa strongly arched, apex rounded, termen very obliquely rounded; leaden-grey, a broad dark fuscous median suffusion from base to costa before apex; in this discal dots are with difficulty discernible, first discal at one-third, plical before it, second discal at two-thirds; some dark fuscous irroration in terminal area; cilia grey. Hindwings grey-whitish; cilia grey-whitish, on apex grey.
Queensland: Charleville, in October; one specimen from larva feeding on joined leaves of Hucalyptus.
302. OENOCHROA SUFFULVA, 0. SD. suffulvus, rather tawny.
9. 18 mm. Head and thorax fuscous, the latter with a small posterior crest. Palpi with second joint slightly expanded at apex, reaching base of antennae, terminal joint three-fifths; fuscous. Antennae fuscous. Abdomen brownish-grey; apices of segments paler. Legs fuscous; posterior pair ochreous-whitish. Fore- wings narrow, costa gently arched, apex rounded, termen very oblique; fuscous- grey; an inwardly oblique whitish streak at about one-third, not reaching either margin, immediately followed by a suffused dark spot; an obscure discal dot at three-fourths; cilia fuscous-grey. Hindwings whitish-grey-brown, towards apex grey; cilia grey.
Characterized by brownish hindwings and oblique mark on forewings.
North Queensland: Stannary Hills near Herberton (Dr. T. Bancroft). Queens- land: Banana, in March (Mrs. Hobler). Two specimens.
305. OENOCHROA ZALOTYPA, N. SD. fadorumos, with stormy markings.
6. 20-23 mm. Head fuscous; face whitish. Palpi with second joint reaching base of antennae, terminal joint three-fifths; ochreous-whitish irrorated with whitish; terminal joint fuscous except extreme apex. Antennae grey; ciliations in male 24. Thorax fuscous. Abdomen grey. Legs fuscous with ochreous-whitish rings; posterior pair ochreous-whitish. Forewings suboblong, costa gently arched, apex round-pointed, termen obliquely rounded; ochreous-whitish densely irrorated with fuscous, and with some brown scales, especially on veins; ill-defined fuscous spots on costa at one-fourth and middle; stigmata dark fuscous, first discal at one-fourth, confluent with plical, which is before it, a pale dot above and beneath, second discal just beyond middle, rather large, an additional dot before and beneath it; a large suffused fuscous subapical spot, from which proceeds an obscure line to tornus; cilia grey with a fuscous median line. Hindwings and cilia whitish-grey. .
W.A.: Denmark in March; four specimens received from Mr. W. B. Barnard, who has the type.
307. OENOCHROA DYSTENA, N. SD. dvaTynvos, miserable. 3, 2. 17-20 mm. Head and thorax grey; the latter with a small posterior crest. Palpi with second joint dilated with rough scales at apex anteriorly,
BY A. J. TURNER. 115
reaching base of antennae; terminal three-fifths; grey, second joint whitish towards base. Antennae grey; ciliations in male 1. Abdomen grey; tuft whitish- grey. Legs dark grey; posterior tibiae ochreous-whitish. Forewings oval, costa moderately arched, apex pointed, termen very oblique; grey; stigmata small, obscure, fuscous, often wholly or partly obsolete; first discal shortly before middle, plical much before it, second discal at two-thirds; cilia grey. Hindwings and cilia grey.
A poor specimen of this obscure species, which I sent to Mr. Meyrick many years ago, was placed by him under O. homora Meyr. Having seen a good example of that species from Sydney, I am now satisfied that this is distinct.
Queensland: Brisbane, in August, September, February and May; eight specimens.
41. Gen. PLAcocosMA, Nn.g.
Meyr., Proc. Linn. Soc. N.S.W., 18838, p. 333. Type, P. anthopetala Meyr.
Tongue present. Palpi- ascending, recurved; second joint reaching base of antennae, thickened with smoothly appressed scales; terminal joint shorter than second, slender, acute. Antennae with basal pecten; in male with moderately long ciliations. Thorax with a small posterior crest. Forewings with 7 to apex. Hindwings with 5 from middle or above middle of cell.
I know only the type species, and would not be sure of the exact position of this small genus.
Three Species: 7311, diantha Meyr., Exot. Micro., i, 1913, p. 134 (Darwin) .— +312, resumptella W1k., xxix, p. 681 (Sydney). = hephaestea Meyr., Proc. Linn. Soc. N.S.W., 1883, p. 333.—313, anthopetala Meyr., ibid., 1883, p. 333 (Brisbane, Sydney).
THE PETROLOGY OF THE HARTLEY DISTRICT. III. THE CONTACT METAMORPHISM OF THE UPPER DEVONIAN (LAMBIAN) SERIES.
By GerMAInE A. JopLin, B.Sc., Department of Mineralogy and Petrology, Cambridge. Junior International Fellow for 1933-34 of the International Federation of University Women.
(Plate i; three Text-figures.)
[Read 27th March, 1935.]
INTRODUCTION AND PREVIOUS RECORDS.
The contact altered sediments of the Hartley district belong to the Lambian Stage (Brown, 1931) of the Upper Devonian Series. There are various records of the contact effects produced by the Hartley—Bathurst bathylith, but a number of these describe altered rocks that belong to a different sedimentary series. C. S. Wilkinson (1877) reported on the occurrence of wollastonite, epidote and garnet in rocks from Kirk’s Farm on the Fish River, near Oberon; and as far as the present writer is aware these are altered Lambian sediments. The earliest record of the Hartley hornfelses themselves is to be found in a note by G. W. Card (1896) in which he describes a series of specimens forwarded to him by Curran, Ball and Rienits (1896). In his report to the Metamorphic Committee of Section C at the Hobart Meeting of the Australasian Association for the Advancement of Science (1928), W. R. Browne gives a later reference to the Hartley hornfelses, and they are again mentioned in the Sydney Handbook of the Australian and New Zealand Association for the Advancement of Science (1932). No detailed work, however, has been done on this aureole, and it is hoped that the present paper will contribute to our knowledge of this large and interesting bathylith.
Herein is given a detailed account of the metamorphism of the Lambian series, which consists of arenaceous, argillaceous and areno-calcareous sediments with intercalated lavas of acid and intermediate character.
SEQUENCE AND STRUCTURE.
It has been found convenient to divide the Lambian Stage, as developed at Hartley, into an upper and a lower series. The upper consists essentially of quartzites and is unfossiliferous. The lower includes all those types that are associated with the fossiliferous quartzites.
The upper series is found capping the spurs of the entrenched Cox’s River and its tributaries. It is directly overlain by the Kamilaroi Upper Marine, and is usually deeply weathered. There appears to be little variation in the rock type, but this may be due to weathering which tends to produce uniformity. Highly jointed quartzites with intercalated “purple-hornfelses” comprise the upper series.
The lower series outcrops along the river and in the beds of the tributary
BY GERMAINE A. JOPLIN. 17
creeks, and good unweathered sections may hence be obtained. Massive quartzites, crowded with Spirifer disjunctus, and interbedded with bands of “purple-hornfels” of varying width, are by far the most prominent beds in this series. Grits passing in places into conglomerates, calcareous cherty rocks and argillaceous types are developed to a lesser extent. Sills of altered porphyrite occur on several different horizons, and it is believed that the felsites outcropping on Cox’s River, on Pine Ridge Creek, and on Moyne Farm, represent a flow at the base of the lower series. As most of the hornfelses with which this paper deals were collected from the lower series, it is pertinent to describe this series in a little further detail.
The Fossiliferous Quartzites and ‘“Purple-hornfelses” are very intimately associated and, as has been stated above, the latter are also prominent in the upper series. With the fossiliferous quartzites, they occur in bands varying from a few millimetres to many feet in thickness. Occasionally they appear to be inter- mingled with the quartzites, and sometimes they predominate altogether and the quartzite occurs only as streaks and blebs in the ‘“‘purple-hornfels’. This latter type is more prominent in the upper part of the lower series, where only occasional fossiliferous quartzites are developed in association with great thicknesses of “purple-hornfels”, which is associated here also with the more argillaceous beds. In the lower part of this series the calcareous quartzite is the predominating type, and the “purple-hornfelses” appear only in comparatively narrow bands. The name “purple-hornfels” is a convenient field term and, though it includes several different types of hornfels, all appear similar in the hand-specimen. They are extremely fine-grained rocks with a subconchoidal fracture, and show a charac- teristic purplish-brown colour, due to the presence of biotite.
In the quartzites the fossils are restricted to comparatively narrow bands. Such bands vary from a few inches to about a foot in thickness, and are crowded with Spirifer disjunctus, with an occasional Rhynchonella pleurodon. Between these richly fossiliferous bands the quartzite is quite barren, but the composition of the hornfels shows that a certain amount of lime was contained in the matrix of the original sediment.
Lithologically the ‘“‘purple-hornfelses” and the fossiliferous quartzites pass imperceptibly into banded calcareous cherts in which the Spirifer bands are still abundant. The calcareous radiolarian cherts described below have never been found in association with the fossiliferous type, though both rocks are developed in the upper part of the lower series. These rocks bear a remarkable resemblance to the banded calcflintas and killas of Cornwall (Ussher et al., 1909; Reid et al., 1910).
The Porphyrites and Banded Cherts also show a close field association. The former usually occur as sills, though on Hughes’ Creek a transgressive relation is apparent. On the map (Plate i) the thickness of these sills has been slightly exaggerated, and in some cases a single wide sill has been shown instead of two narrower ones separated by cherts, as on Liddleton Creek and Moyne Creek (see Text-fig. 1).
The cherts are usually distinctly banded, and Dr. W. R. Browne has found traces of radiolaria in the Moyne Creek type. On Cox’s River, just above Marriott’s Creek, a slightly different type of chert occurs. This is a pale pink rock with numerous dark purplish bands and blebs suggesting the ‘“purple- hornfels” and its associate. No fossils, however, have been found in this rock, but the composition of the hornfels indicates a calcareous silt.
E
18 PETROLOGY OF THE HARTLEY DISTRICT. iii,
GEOLOGICAL SKETCH MAP
On) Eve
MOYNE ANTICLINE-
> Basic Dyke Granite Quartz-mica-diorite .
Quartzite
“Purple - Hornfels’ elc.
‘ By Altered Porphyrite
FES |Cole-slicat Hornfels .
UPPER DEVONIAN
t 6 Wi ye 7 ey Z Ley ye2 A ZR Kl PER Ao NAmbhible X % X xX of a Plagioclase = Biotite Hornfels
Text-fig. 1.
BY GERMAINE A. JOPLIN. 19
With regard to the structure of the Devonian beds, it has already been stated (Joplin, 1933) that they occupy a broad syncline between the northern and southern outcrops of granite. The major fold, which trends approximately east and west and pitches to the east, is turned over abruptly into sharp anticlines at the granite contacts. The basic stocks of the plutonic complex (Joplin, 1931, 1933) are injected into the trough of the syncline, and there is reason to believe that they are associated with fault zones.
The exact nature of the felsites on Cox’s River and on Pine Ridge Creek is doubtful. They may represent pre-granite intrusions of an irregular nature, which have been subsequently contact-altered by the granite; or, what seems more likely, they may represent a basal flow which has been duplicated by faulting.
Reference to Plate i will show that the sequence of the beds indicates a repetition which is suggestive of a north-south fault along the western boundary of the eastern felsite. Actually there are indications of brecciation both in the felsite and in the adjacent calcareous beds along this margin, and the felsites are very highly jointed. Minor faults have been observed on the north-eastern margin of the felsite; and the displacement of the anticlinal axis is again suggestive of a north-south fault with a throw to the west.
It is difficult to correlate the Moyne Creek section without postulating another fault of considerable magnitude. The close association of the porphyrites and cherts on Moyne Creek and Liddleton Creek suggests a datum horizon. The occurrence of a small patch of felsite on Moyne Farm is another difficulty that might be explained by faulting.
It is not improbable that complex faulting would occur along a contact where there have been successive periods of injection. Only detailed mapping and contouring of this fairly rugged area will show the extent of such faulting and, as far as the present study is concerned, there is little to be gained from a piece of work that would involve such an expenditure of time. Nevertheless, the writer has pointed out (1933) that the nature of the Cox’s River Intrusion is suggestive of faulting in that area, and it seems not impossible that both the basic stocks have been associated with fault zones. Moreover, if no faulting be postulated, there would be an unbroken succession of Lambian strata over a distance of about four miles from east to west, and as the dip is usually at a high angle, this would give an abnormal thickness. On the other hand, if faults be assumed to be present west of the river and west of Moyne Creek, an approximate estimate gives a thickness of about 2,000 feet, which is in accord with more recent observations at Rydal.
On Plate i certain of the arrows indicating dips do not show the amount of dip. Where the amount of dip is indicated, measurements have been made with a clinometer rule, and where such is not shown, general compass directions have been taken along the strike of the bed.
CORRELATION WITH THE Type SECTION AT Mr. Lamsiz, RypDAL.
In 1896 G. W. Card pointed out that the Hartley sediments “may be regarded as the eastern extension of the Mt. Lambie Beds”. In the Rydal district, as at Hartley, Spirifer disjunctus occurs abundantly in restricted bands in a massive quartzite. On Mt. Lambie these beds are apparently unmetamorphosed and this prompted the writer to look for the bands corresponding to the Hartley ‘“purple- hornfelses”. They were found to be represented by soft reddish-purple shaly rocks which readily weather away, and appear quite insignificant among the resistant
20 PETROLOGY OF THE HARTLEY DISTRICT. iii,
quartzites. That the “purple-hornfelses”, in their unaltered condition, correspond to the so-called ‘‘red shales” is further supported by the presence of an occasional pebble of “red shale” in the Kamilaroi conglomerate at Hartley. It is possible that these soft red rocks represent either fine-grained periodically extruded tuffs, or fine silts brought down by floods. A microscope examination of the Rydal rock shows small angular fragments of quartz and lends support to the former view. This material, whatever be its origin, has possibly been responsible for the sudden periodic killing off of the Spirifers, which evidently formed massive shell banks along the shallow coast.
' At Rydal the Spirifer beds pass up into grits, “red shales’, buff shales and quartzites and, if the so-called “red shales” may be taken as the equivalents of the ‘‘purple-hornfelses”’, the sequence at Rydal closely corresponds to that at Hartley.
To the west of Mt. Lambie, below the Spirifer beds, there is an igneous rock, which may correspond to the basal felsite at Hartley.
As far as the present writer is aware, there are no porphyrites in the Rydal district, and only one bed of calcareous chert is known. This occurs near the top of the series just to the west of Rydal railway station. It is probable that the sills and their associated cherts are developed quite locally at Hartley.
At Rydal the general direction of strike is north and south, whilst at Hartley (10 miles distant from Rydal) the axes of the folds trend approximately east and west. It is possible that the intrusive masses of granite to the north and south of Cox’s River at Hartley have acted as the jaws of a vice in which the sediments have been squeezed into their present position.
WIDTH OF THE ConTAcT AUREOLE.
There are three difficulties in the way of measuring the width of the contact aureole, and of zoning the progressive changes as the igneous boundary is approached: (1) Owing to the close proximity of the overlying Kamilaroi strata, no unaltered Devonian rocks are exposed, and there is thus no standard of comparison; (2) the contacts of the basic stocks and of the granite are so close that if a bed be traced out of the aureole of one intrusion it immediately enters that of another; (3) apophyses some distance from the apparent boundary, as well as large inclusions of sediments in the igneous rocks, suggest that the roof of the bathylith has not been completely removed; there is reason to believe, therefore, that the gradient of the intrusions is fairly shallow, and thus linear distances measured from the apparent contact are obviously incorrect. For reasons stated above, therefore, very little information may be gained by tracing a single bed along its strike, but in the next section it will be shown that some idea of the intensity of metamorphism, and of the width of the inner zone of hornfelses, may be gained by an examination of a series of specimens of the same rock type from different parts of the area.
The first of these difficulties may be overcome to some extent by a comparison with unaltered rocks at Rydal.
In the petrographical sections, distances from the apparent contact are always stated, but it must be borne in mind that these are not necessarily correct, and that the actual contact may be much closer. The section dealing with incipient metamorphism indicates some of the anomalies that arise, if this be disregarded.
In a general way, it may be stated that the contact is widest in the arenaceous, areno-calcareous and calcareous chert beds, and less wide in those that contain
BY GERMAINE A. JOPLIN. 21
-an appreciable amount of shaly material. Thus it is shown that calc-silicates, such as diopside, amphiboles and epidote, may develop as well-formed minerals, when associated more argillaceous rocks show only an incipient development of biotite.
As it seems very evident that distances cannot be measured from the true contact, it is useless to give figures for the width of the inner zone of hornfelses.
INCIPIENT METAMORPHISM.
It has been shown above that difficulties attend the study of this phenomenon, and the present section deals with a description of unmetamorphosed specimens from Rydal, and of scattered rocks at the greatest possible distance from the contact at Hartley. The main rock types that have given rise to the hornfelses in the Hartley district are: (i) “red shales”, (ii) fossiliferous quartzites, (iii) calcareous cherts, (iv) sandstones and grits, (v) normal shales.
(i) Two of the so-called “red-shales” from Rydal have been examined, and are found to consist of small angular chips of quartz and a little alkaline felspar set in a matrix of chlorite with a small quantity of white mica. Greenish biotite, magnetite, sphene, zircon and tourmaline are accessories. The chlorite is much stained by haematite, which gives the rock its red colour.
In the Hartley region a rock in Deep Ravine, at a distance of 660 yards from the apparent contact, shows some evidence of metamorphism. It is exactly similar to the “purple-hornfelses” in the hand-specimen, but under the microscope a slightly clastic structure is apparent and the rock consists of quartz and alkaline felspar grains surrounded by a matrix of tiny flakes of greenish-brown mica and a little chlorite. This rock is something of an anomaly, and the contact is possibly closer than is apparent. The typical reddish-brown authigenic biotite has been noted at a distance of about 450 yards from the contact, and incipient brown biotite enters at 580 yards.
(ii) Fossiliferous quartzites from Mt. Lambie and from Solitary Creek, Rydal, have been examined, and in both cases calcite is conspicuously absent. Occasion- ally groups of calcite crystals have been noted in the field, and it appears that the carbonates have been removed by leaching. Though unaffected by contact meta- morphism, the Rydal quartzites show evidence of silicification, which is possibly due to cementation (Van Hise, 1904), and it would appear that the lime had been removed during this process. In the hornfelsed type, where lime is fixed in the form of a silicate, it may be preserved. Both quartzites consist of quartz and a little alkaline felspar in a matrix of chlorite. Accessories are white mica, sphene, magnetite and haematite. It is believed that the fossiliferous quartzites at Hartley originally had a composition rather similar to this, and that calcite was present in the matrix as well.
At Hartley the fossiliferous quartzites do not occur at a greater distance than 580 yards from the contact, and at this distance the effects of thermal meta- morphism are apparent. Hand-specimens show well preserved fossils and ‘‘nests’”’ of secondary calcite crystals.
Under the microscope the rocks still show their clastic structure, but the fine-grained groundmass is entirely recrystallized and consists of quartz, basic plagioclase, diopside, amphibole and sphene. Small patches of calcite are also present, and though they appear to have been recrystallized, the temperature has not been sufficiently high for the formation of wollastonite. Wollastonite occurs abundantly at the actual contact in several localities and has never been found at a greater distance thdn 350 yards from the apparent contact. In most
22 PETROLOGY OF THE HARTLEY DISTRICT. iii,
of these cases wollastonite may be seen replacing the actual fossil, and recognizable Spirifers, partly changed to wollastonite, have been collected within a few inches of the contact. Shells pseudomorphed by aggregates of diopside, sometimes containing a little epidote or amphibole, have been noted at 580 yards, and the associated “purple—hornfels” bands show a development of incipient biotite.
(iii) One example of calcareous chert has been collected in the Rydal district, and it is quite unaffected by thermal metamorphism. It is a very fine-grained rock consisting mainly of quartz with a matrix of chlorite and a little calcite. Magnetite, biotite and a little plagioclase are also present, and zircon occurs as an accessory. No specimen of this rock has been found outside the inner zone of hornfelses at Hartley.
(iv) A rock occurring on top of the ridge between Deep Ravine and Bonnie Blink Creek lies at a distance of about 850 yards from the granite, but in the hand-specimen it is a fairly typical quartzite.
Under the microscope, however, there is a distinctly clastic structure apparent. Large (0-4 mm.) somewhat rounded grains of quartz and alkaline felspar are surrounded by a matrix of sericite and a little chlorite, and minute flakes of incipient biotite are just discernible. Magnetite and zircon are accessories.
(v) No unaltered or partly altered normal shales are known.
From these scanty observations it would appear that the width of the contact varied in the different beds and that the calcarous rocks responded to the thermal effects before the more argillaceous types.
In the more porous sandstone hornfelses incipient biotite is noted at 850 yards, but in the more compact “‘red-shales” it does not make its appearance until within 580 yards of the contact.
PETROGRAPHY OF THE HORNFELSES OF THE INNER ZONE. (i) Andalusite-cordierite-biotite Hornfelses.
Three examples of this class have been recorded from different parts of the aureole. They are fine-grained, dense, dark grey rocks. One, near the road crossing on the southern branch of Grant’s Creek, at a distance of 880 yards from the granite and 660 yards from the diorite, shows a faint spotting, which under the microscope is seen to be due to aggregates of quartz grains associated with andalusite and flakes of muscovite. ;
A rock from Moyne Creek, at a distance of 130 yards from the granite and 390 yards from the diorite, is fairly typical of this class. Its structure is grano- blastic with an average grainsize of about 0:15 mm. The constituent minerals are quartz, andalusite, altered cordierite, biotite, orthoclase, magnetite and a little muscovite, chlorite and tourmaline. Rutile and sphene have been noted as accessories in rocks of this class.
The andalusite occurs in small stumpy prisms (averaging 0-1 mm.), which often show a strongly pleochroic rose-pink core. There is a slight marginal alteration to sericite. The cordierite is entirely altered into a green micaceous substance, and occurs in large aggregates of ill-formed stumpy prisms, or more commonly as xenoblasts. Small flakes of biotite are associated with these pseudo- morphs and probably represent inclusions in the original cordierite. Deep-brown biotite (a’ = 1:592, 7’ = 1-637) oceurs in poikiloblastic flakes measuring up to 0-3 mm., and, though more frequently associated with the cordierite areas, is present to a lesser extent in the andalusite-quartz and andalusite-quartz-orthoclase areas.
BY GERMAINE A. JOPLIN. 23
A partial analysis of this rock is shown in Column I below:
1 Il. SiO, do Ree ae 616 O10 50 67-00 62-80 Al,03 a. ay a Me a oe BEY 19°74 Fe,03 a is o ae on si 0°75 0:00 FeO 4-34 1-98 MgO 1-16 1:34 CaO aie ee aD oo So ae 0:66 0:87 Na,O 5% se ah Os sie oe 2-14. 22 K.O He 5-44 6-56 H,0+ boll meen 0-27 H,0 — CAs 0-86 + Loss on Ignition, C bi0 So _— 1-58 PROWL TiO,.. 0:03 1-36 P,0; abs 0-60 MnO pnd 0:02 Stns nd. 0-52 100-24 99-72 Less O=S 0:23
I. Andalusite-cordierite Hornfels, Moyne Creek, Por. 124, Parish of Hartley. Anal. G. A. Joplin.
Il. Andalusite-cordierite Hornfels (Class 1), Gunildrud, Contact of Soda-Granite, Christiania. Anal. M. Dittrich. V. M. Goldschmidt, Die Kontaktmetamorphose im Kristianiagebiet. Videnskap. Skrift. I. Math.-Nat. Kl., No. 1, p. 148, 1911.
It appears that the main difference between these rocks lies in the greater abundance of andalusite and orthoclase in the Christiania hornfels, and the excess of quartz, biotite and magnetite in the Hartley rock.
In the field this rock is closely associated with a rather mottled, lighter grey hornfels. Under the microscope these are essentially the same, but the latter contains in addition an abundance of white mica and tourmaline. The biotite is also somewhat altered to chlorite. A very similar type of hornfels occurs on Cox’s River below the mouth of Marriott’s Creek at a distance of 400 yards from the diorite.
A very much altered rock is met with on Bonnie Blink Creek, and it is possible that it may belong to this class. In the hand-specimen it is a banded grey hornfels with rows of black rectangular spots which consist entirely of sericite and muscovite. Remnants of cordierite have been recognized, and it is possible that the dark spots were originally andalusite.
(ii) Andalusite-biotite-orthoclase Hornfels.
A rather unique type has been collected as a boulder in Bonnie Blink Creek. It is light purplish-grey rock containing abundant pinkish-white spots which measure about 6 mm. and stand out in relief on weathered surfaces. These spots are prismatic crystals of andalusite which show a good deal of sericitization.
The fine-grained groundmass consists of orthoclase, very abundant reddish- brown authigenic biotite (a’ = 1:595, 7’ = 1-635), muscovite and a little quartz. Accessory minerals are greenish zircon, magnetite, tourmaline and sericite. The zircons commonly occur as inclusions in the andalusite.
(iii) Cordierite-quartz Hornfelses. C. E. Tilley (1924) has divided these hornfelses into (a) Biotite-rich and (b) Biotite-free types. At Hartley no hornfels of the type absolutely free from biotite
24 PETROLOGY OF THE HARTLEY DISTRICT. ili,
has been recorded, but a number contain such a small amount of this mineral that they stand out in marked contrast to the Biotite-rich division, and it is proposed to consider them separately as Biotite-poor types.
(a) Biotite-rich Types—These hornfelses are developed abundantly on Cox’s River and Bonnie Blink Creek, and one example has been collected from the contact on Yorkey’s Creek. Except for the total absence of plagioclase these rocks are similar to the cordierite-plagioclase assemblage described below. The constituent minerals are cordierite, quartz, biotite, orthoclase, magnetite and a little white mica. Accessory minerals are zircon and apatite, and a little sphene has been noted in a few examples. As in the more calcareous type described below, cordierite may occur as oval porphyroblasts giving the rock a spotted appearance, or it may form small xenoblasts in an even-grained granoblastic rock.
A typical example of the even-grained type occurs on the spur between the river and the junction of Liddleton and Bonnie Blink Creeks. It is a rather coarse-grained, resinous, greyish-brown rock, which, on weathered surfaces, shows a distinct banding. Under the microscope several types of banding may be recognized—differences in grainsize, alternations of biotite-rich and biotite-poor types, cordierite-rich seams and selectively altered cordierite seams.
In most of these rocks cordierite is very abundant, and several good examples of twinning have been noted. In longitudinal section multiple twinning is apparent and in cross section the mineral breaks up into sectors. The cordierite is frequently altered both to aggregates of white mica and to yellow, isotropic pinite. In some of the cordierite-rich seams this mineral is clouded by minute inclusions of iron ore. These evidently represent iron-rich chlorite seams in the original sediment.
(6) Biotite-poor Types.—It is stated above that these hornfelses occur inter- bedded with a biotite-rich assemblage near Cox’s River. Another example occurs on Moyne Creek. Except for a marked decrease in biotite, a concomitant increase in orthoclase and magnetite and the total absence of white mica, these rocks are very similar to the above and need no further description.
The table below is an analysis of a cordierite-quartz hornfels containing a small amount of biotite; it is regarded as fairly typical of this class of hornfels. Itisa medium-grained granoblastic rock consisting of cordierite, quartz, orthoclase, magnetite, and a little biotite (a’ = 1-587, B’ = 1-630, y’ = 1-633) and white mica.
The analysis used by C. E. Tilley (1924) in his discussion on this class of hornfels has been included to show that the resulting mineral assemblage is independent of the amount of quartz in the original rock. It is evident that the Hartley rock was a sandstone with an iron-chlorite matrix, whilst the rock in Column II represents an original chlorite-rich shale. As pointed out by Prof. Tilley, the mineral assemblage in these hornfelses depends upon the RO/R,.O, ratio.
(iv) Cordierite-plagioclase Hornfelses.
A number of examples of this type are recorded from the contacts on Moyne and Grant’s Creeks and, with but two exceptions, they occur within 5 yards of the igneous boundary. One rock of this type is found on Grant’s Creek at a distance of 220 yards from the diorite and apparently 400 yards from the granite, but the fact that it is invaded by veins of tourmaline-aplite suggests an underground extension of the granite. Another example is recorded from near the head of Horse Hole Gully at a distance of 700 yards from the diorite.
BY GERMAINE A. JOPLIN. 25
I Hite SiO, 84:23 59-83 Al.O; Geos 17:47 Fe.0, 2-06 4-09 FeO 1-61 3°93 MgO 0:64 3°70 CaO 0:78 0:49 Na,O 1-20 1:08 K,0 1:53 4°42 H,O 0-57 3°80 TiO, tr: 0:93
MnO or = P.O; abs 0:18 SO, . — 0-13 99-75 100-05
I. Cordierite-quartz-biotite Hornfels, from granite contact on hillside above junction of Liddleton and Bonnie Blink Creeks, Por. 27, Parish of Lowther. Anal. G. A. Joplin.
Il. Cordierite-quartz-biotite Hornfels, Abbenstein (Harz), described by O. H. Erdmannsdorffer (Jahrb. Preuss. Geol. Landesanst., Vol. xxx, 1909, p. 357). Quoted by C. E. Tilley (Quart. Journ. Geol. Soc., 1924, p. 37).
These rocks fall into three groups—a spotted type, a massive resinous dark grey hornfels, and a type very rich in biotite with indications of a parallel structure.
The rock on Grant’s Creek, Por. 124, Par. of Hartley, is a typical spotted hornfels. It is a dense, dark purplish-grey rock crowded with resinous, black oval spots about 2 mm. in length. On weathered surfaces pitting is conspicuous.
Under the microscope the hornfels is seen to consist of numerous oval porphyroblasts of cordierite set in a fine granoblastic groundmass of biotite, quartz, plagioclase and orthoclase. Accessory minerals are magnetite, tourmaline and zircon. The cordierite is extremely fresh, and is crowded with inclusions of pale greenish-brown biotite, quartz and plagioclase. The biotite inclusions are by far the most abundant, and are of a paler colour than the biotite of the groundmass. In the groundmass flakes of biotite are particularly abundant as a fringe around the porphyroblasts, and this is probably due to the throwing out of inclusions during advancing metamorphism. The biotite of the groundmass occurs in humerous, strongly pleochroic, reddish-brown flakes (a’ = 1-587, 6’ = 1-627, 7 = 1:633). The colour, pleochroism and refractive indices indicate a high iron content. Plagioclase is sometimes twinned and appears to be andesine.
The rock exhibits a slight parallelism due to the arrangement of the biotite flakes and of the longer axes of the porphyroblasts. This appears to be the original direction of bedding.
The tourmaline has no doubt been introduced by the tourmaline-aplite that invades the hornfels. In this rock muscovite is absent.
The analysis of this rock is given in Column I below, where it is compared with analyses of similar assemblages cited by Goldschmidt (1911). Except for a slightly greater abundance of silica and lime, and a little less magnesia, the
26 PETROLOGY OF THE HARTLEY DISTRICT. iii,
Hartley rock is intermediate in composition between these hornfelses. A strict comparison made on the basis of specific gravity might indicate closer affinities.
de Tite III. SiO, .. ae “es 61:50 58°83 56-88 Al,O; .. 19-84 17-54 20-68 Fe.0, 1-39 0-00 2-66 FeO 5-20 8-42 4°54 MgO 2-67 3-40 3-15 CaOlier 2°91 2-24 1-29 Na,O .. 1-11 1-35 0-91 LEO) ge 4-39 4-35 7-49 H,0+ 1-28 1-96 SD one H,0 — 0-04 0-13 i TiO, Mi ap 0-42 0-59 — MnO .. 3 e. tr. 0-09 — 1240)3. 6 a ae abs. 0-46 — Ci) ieee S6 se = 0-50 — 100-75 99-86 100-12
I. Cordierite-plagioclase Hornfels, Grant’s Creek, Por. 124, Parish of Hartley. Anal. G. A. Joplin.
II. Cordierite-plagioclase Hornfels (Class 3), Kolaas, contact of the nordmarkite, Christiania. Anal. M. Dittrich. V. M. Goldschmidt, Die Kontaktmetamorphose im Kristianiagebiet. Videnskap. Skrift. I. Math.-Nat. Kl., No. 1, 1911, p. 156.
III. Cordierite-plagioclase Hornfels,-Monte Doja, Adamello. Pelikan (Tscher. Min. Pet. Mitt., 12, 1891, p. 156). Quoted by Goldschmidt. TIbid., p. 157.
An example of the massive, resinous type of hornfels occurs on Grant’s Creek just above its junction with Moyne Creek at about 1 yard from the contact. It is a granoblastic rock with slightly coarser grainsize (0-6 mm.), and contains the same mineral assemblage as above. The biotite is less abundant, and is of a more reddish colour with R.I. a’ = 1-588, p’ = 1-635, y’ = 1-637. Cordierite is represented by masses of secondary mica.
The biotite-rich members of this class occur at the mouth of Moyne Creek and are banded with biotite-plagioclase and biotite-amphibole-plagioclase assem- blages. The biotite may be arranged in criss-cross fashion, but is more often parallel to the original bedding.
(v) Plagioclase-biotite-quartz and Biotite-quartz Hornfelses.
These types are perhaps the most widely distributed in the Hartley aureole and represent the largest bulk of the “purple-hornfelses”’.
A hornfels of this type occurs in the upper series at the top of the spur to the west of Grant’s Creek, Por. 118, Parish of Hartley, at a distance of 200 yards from the granite. Under the microscope it is seen to consist of a fine mosaic of quartz, biotite, plagioclase and orthoclase, with tourmaline in large irregular aggregates. There are very small veins of igneous material associated.
A typical example of this hornfels, containing a small amount of plagioclase, has been analysed (see Column I below). It occurs on Bonnie Blink Creek at a distance of 440 yards from the granite. It is a fine granoblastic rock consisting
BY GERMAINE A. JOPLIN. 27
of quartz, biotite (a’ = 1:585, 7’ = 1-633), orthoclase, plagioclase, ilmenite and accessory zircon.
I IL. III SiO, 82:27 79-28 47-93 NWO, « « 9-32 6-60 20-34 Fe,0; abs. 0-51 4°35 He Omer 2-65 2-32 8-63 MgO .. 1-09 1-96 5-58 CaOnee 1-80 3°95 1-64 Na,O .. 0-83 3°27 4°70 K.0 1:27 0-96 4°88 H.0 0-72 0-72 0-72 TiO eee as b, 0-47 0-40 0-76 MnO .. be et nd. 0-25 0-13
PlOn, * aes abs. 0-1 =
COP: # ¥ = 0-09 ae 100-42 100-42 99-66
I. Biotite-plagioclase Hornfels (‘‘Purple-hornfels’”), Bonnie Blink Creek, Little Hartley. Anal. G. A. Joplin.
Il. Felspathic Hornstone of the Cale-flinta Series, Tregullan, 14 m. SSW. of Bodmin, Cornwall (Slide E5458). Anal. H. G. Radley. W. A. Ussher et al., Mem. Geol. Surv. Hung. and Wales, Sheet 347, 1909, p. 101.
III. Biotite-plagioclase Hornfels (Class 3), Christiania. Anal. M. Dittrich. V. M. Goldschmidt, l.e., 1911, p. 37.
The Cornish hornfels occurs associated with calc-flintas, as does the one from Hartley. The Hartley rock appears to be less rich in plagioclase, but biotite and orthoclase are possibly more abundant. The Christiania hornfels has been included for contrast. This again emphasizes the fact that a similar mineral assemblage may arise in a shale or in a siliceous rock with a shaly matrix. There are other rocks of a very similar appearance in which plagioclase cannot be identified, and it is believed that these represent lime-poor assemblages related to those described above.
It will. be shown later that with an increase of lime and magnesia these rocks pass into amphibole-bearing types from which they cannot be distinguished in the hand-specimen.
One example of the biotite-plagioclase assemblage occurs at the mouth of Moyne Creek, where it is interbedded with cordierite-plagioclase-biotite and amphibole-plagioclase-biotite types. All three types are much coarser in grain- size than the “purple-hornfelses’, which they closely resemble in mineral constitution, and their origin will be discussed later. They are often veined with igneous material.
(vi) Amphibole-plagioclase-biotite Hornfelses. These rocks have been collected from within a few yards of the granite near the mouth of Moyne Creek, and from among the ‘cherts” on the northern limb of the Moyne anticline at a distance of 140 yards from the diorite.
28 PETROLOGY OF THE HARTLEY DISTRICT. iii,
The rock occurring at the mouth of the creek has been referred to above; it has a fairly coarse grainsize, is very rich in biotite, and exhibits a parallel structure similar to the associated assemblages which have already been described. The other example is typically a “‘purple-hornfels” in the hand-specimen. Under the microscope the coarser grained rock is seen to consist of biotite, plagioclase, amphibole, orthoclase, quartz, sphene and a little magnetite and/or ilmenite, tourmaline and pyrites.
The amphibole forms highly poikiloblastic plates (0-5 mm.) which are arranged in linear fashion, and evidently represent calcareous seams in the original sediment. The amphibole is green, markedly pleochroic, with an extinction angle of about 22°, and R.I. a’ = 1:637, 7’ = 1-658. It is optically negative, and is thus a common hornblende near pargasite. The biotite shows a parallel arrangement which is in the same direction as the strings of amphibole xenoblasts. It is a strongly pleochroic reddish-brown type with R.I. a’ = 1-580, 7 = 1:633. The plagioclase is frequently twinned and occurs in small xenoblasts (0:1 mm.). It is andesine (Ab,,An,,) with R.I. a’ = 1-550, y’ = 1-555.
Another rock of this type, also from Moyne Creek, is a little more calcareous and contains a nodule consisting almost entirely of large (3 mm.) sub-idioblastic crystals of amphibole, with refractive indices a’ = 1-616, 6’ = 1-625, y’ = 1-635, and an extinction of about 15°. Thus, according to Winchell (1933), the mineral belongs to the tremolite-pargasite series, and has a composition Tr,;Pr,;.
(vii) Amphibole-diopside-plagioclase-biotite Hornfelses.
Only a few examples of this type have been recorded from the aureole. They occur on Moyne Creek, Bonnie Blink Creek, and on the river just above the mouth of Marriott’s Creek.
Except for the entrance of a little granular diopside these rocks are essentially the same as the fine-grained types referred to above, and, like them, they occur among the so-called ‘‘cherts”.
(viii) Amphibole-diopside-plagioclase Hornfelses.
These are usually fine-grained rocks constituting some of the lighter bands in the calcareous cherts.
One coarse-grained example occurs on Moyne Creek at the contact of a large granite apophysis. It is a mottled light and dark greenish-grey rock which, under the microscope, is seen to consist of large (0-75 mm.) highly poikiloblastic sheets of amphibole and smaller granules of diopside in a groundmass of plagio- clase, quartz and orthoclase, with accessory sphene, zircon and magnetite. A little epidote and clinozoisite and a few flakes of biotite are also present. Scattered hexagonal pseudomorphs consisting mainly of chlorite and clinozoisite possibly represent cross-sections of biotite. The amphibole has an extinction of 20°, and the refractive indices (a’ = 1:635, y’ = 1:655) and optically positive character indicate pargasite near common hornblende. Large pleochroic haloes are frequent around inclusions of zircon.
A coarser more quartzose member of this class occurs on the hillside north- west of the junction of Liddleton and Bonnie Blink Creeks. It contains abundant hollow crystals of pyrites, which are filled with sphene bordered by clinozoisite. A biotite-bearing assemblage is associated.
A rock on Moyne Creek shows this assemblage alternating with seams very rich in magnetite and containing a little biotite.
BY GERMAINE A. JOPLIN. 29
(ix) Diopside-plagioclase Hornfelses. Banding is very common in rocks of this type. It may be caused by alterna- tions with biotite-plagioclase or amphibole-bearing assemblages, by differences in texture and/or by seams consisting almost entirely of pyroxene.
Some of the bands are extremely narrow, and in one slide 1% inches across as many as twelve such alternations have been counted. The pyroxene in some of these banded rocks is of a deep green colour and may contain up to 60% of the hedenbergite molecule (a’ = 1:710, y’ = 1-732). The sharp banding, however, does not admit of an addition of iron from the magma, and this pyroxene possibly arose from layers rich in ferriferous chlorite and calcite, or from mixtures of these minerals with iron ores. Banding in these rocks appears to be indicative of slight fluctuations in sedimentation.
Spotted rocks containing small ellipsoidal aggregates of diopside and plagio- clase, or groups of larger crystals of diopside, are also common in this class. A fine-grained massive type consisting almost entirely of diopside also frequently occurs. Dr. A. Harker (1904, 1932) records cherty diopside-rocks from Skye, where they occur as narrow bands in dolomitic limestones.
All the rocks belonging to this class are chert-like in the hand-specimen, and have a high specific gravity owing to their large content of diopside. They are light-coloured—white, pale pink, grey or, most frequently, pale green. They are extremely like the calc-flintas of the south-west of England.
A typical example from Delaney’s Creek at a distance of 130 yards from the contact may here be described. Under the microscope it is seen to be a fine- grained granoblastic rock with some coarser patches. The constituent minerals are diopside, plagioclase, orthoclase, quartz, sphene, and a little iron-ore. The diopside is very abundant and forms small granules and sub-idioblastic prisms distributed throughout the rock. In the coarser patches the crystals are larger and are always sub-idioblastic. According to Winchell (1933), the composition is) Dizee, (ao = 1-695, 77 = 1-712) and ZAC = 41°. The plagioclase occurs: in extremely minute grains associated with orthoclase. Smal! light-coloured oval patches are numerous and, under strong magnification, are found to consist of diablastic intergrowths of plagioclase and orthoclase. The plagioclase is untwinned and the refractive index is well above that of quartz, but the exact composition cannot be determined. In one of the banded types of slightly coarser grainsize, it has been determined as Ab,An, (a’ = 1:559, B’ = 1:564, 7’ = 1-568).
Under this class might be mentioned a rather unique assemblage consisting of plagioclase, sphene and quartz. The rock in which this type occurs was found as a boulder in the river just below the mouth of Campbell’s Creek. It is a banded rock of the calcareous chert type. It consists mainly of fine “purple- hornfels” with a white calcareous band an inch in width. In the centre of this band there is a seam consisting only of pale green diopside (Di,,He..; a’ = 1-684, 7’ = 1:710) and on both sides of this the sphene-plagioclase assemblage occurs. It would appear that magnesia had been withdrawn from the outer part of the calcareous band and deposited in the central seam, a change which probably took place before metamorphism (see p. 47). In the absence of magnesia the available lime has combined with titania and silica to give sphene. The titania may have been derived from either detrital rutile or ilmenite.
30 PETROLOGY OF THE HARTLEY DISTRICT. iii,
(x) Plagioclase-diopside-epidote Hornfels.
Only one example of this class is recorded from the Hartley aureole. It occurs interbedded with a vesuvianite assemblage and an unstable wollastonite- plagioclase assemblage on the northern limb of the Moyne anticline.
In this rock there seems little doubt that the epidote has arisen as a mineral of primary metamorphic crystallization, and not as a product of metasomatism. The constituent minerals are plagioclase, diopside, epidote, sphene and a little quartz and iron-ore. Orthoclase has not been detected with certainty. The plagioclase forms large poikiloblastic plates or granoblastic aggregates up to 3 mm. across, and encloses granules of epidote and diopside. 'The composition is basic labradorite. Diopside sometimes forms sub-idioblastic crystals 0-4 mm. across, but is more often developed as small xenoblasts intimately intergrown with epidote to form a granular mosaic. In places epidote shows <alteration into clinozoisite and haematite. There is a slight textural banding.
(xi) Plagioclase-diopside-wollastonite Hornfelses.
This is essentially an unstable assemblage, but is by no means uncommon in at least two other aureoles—Deeside (Hutchison, 1933) and Carlingford (Osborne, 1932). A number of wollastonite-diopside rocks containing a very small quantity of plagioclase and scapolite have been collected from many parts of the contact, but there are two examples containing a slightly greater amount of plagioclase, and these will be described here.
One occurs at the diorite contact on Cox’s River just below the mouth of Deep Ravine and, except for the presence of a little andesine, is similar to other rocks with which it is interbedded. It is a cherty green hornfels with bands of radiating wollastonite, which are parallel to the stratification. Under the micro- scope the green cherty layers are found to be made up of a fine aggregate (0-1 mm.) of quartz, diopside, orthoclase, andesine, wollastonite and sphene.
The second example occurs on Moyne Creek, where it is associated with a plagioclase-diopside-epidote assemblage. This rock is also banded. Some of the bands consist almost entirely of andesine with a little wollastonite and diopside. These are intercalated with bands much richer in wollastonite, and which also contain diopside, scapolite, quartz, orthoclase and sphene.
As the Cox’s River rock contains andradite as well, and because the scapolite of the Moyne Creek type is considered to be secondary, both these rocks will be referred to below under the heading of metasomatism.
(xii) Diopside-grossular-wollastonite Hornfels.
Only one example of this hornfels has been recorded. It was collected from the granite contact near the head of Liddleton Creek, and is here associated with wollastonite-diopside assemblages which show extensive metasomatism. In the hand-specimen this rock is green and chert-like, with large masses of radiating, silky wollastonite associated with brown garnet. On the weathered surfaces the wollastonite shows alteration into a white chalky substance described on p. 31. Under the microscope large plates of wollastonite, often measuring more than 6 mm., are seen to be surrounded by or partly wrapped by xencblasts of garnet up to 5 mm. across. Both these minerals contain inclusions of, and are set in a granular mosaic (about 0-1 mm.) of, diopside, epidote, orthoclase, quartz and accessory sphene. In some cases the orthoclase forms large plates, but it is possible that some of this is of igneous origin. The wollastonite may
BY GERMAINE A. JOPLIN. 31
show twin lamellae, and cracks are often filled with calcite. The garnet is apparently of two different varieties—grossular and andradite. The grossular is pale yellow and xenoblastic and is obviously the primary and original mineral in this assemblage. The andradite is deep brownish-yellow and occurs in sub- idioblasts and xenoblasts intimately associated with the lighter coloured variety. Veins of the darker garnet cut through the rock and “blotching” is similar to that observed by G. D. Osborne (1932) at Carlingford. The grossular is some- times anisotropic. Both garnets are poikiloblastic and exhibit a sieve-structure. They frequently form a granular intergrowth with epidote. Like the andradite, much of the epidote is possibly secondary, and this rock will be discussed later in dealing with metasomatism.
(xiii) Vesuvianite-diopside Hornfels.
Only one example of this hornfels has been met with in the Hartley aureole. It occurs as a well-marked band in the centre of the calcareous bed on the northern limb of the Moyne Creek anticline, and is there associated with a wollastonite-orthoclase-diopside assemblage. In the hand-specimen it is a fine- grained chert-like rock showing pale green, white and light brown bands. The rock is highly metasomatized, and it is rather difficult to recognize the original hornfels. It is possible that metasomatism closely followed the recrystallization of the sediment, and that normal thermal metamorphism passed into metasomatism as one continuous process.
As far as can be made out, the hornfels first consisted of a vesuvianite- diopside assemblage in which orthoclase was abundant. Garnet may have been present also. The vesuvianite forms large xenoblasts about 4 mm. across, and often encloses crystals (1 mm.) of altered orthoclase. Orthoclase also forms large independent xenoblasts with a sieve-structure and these, together with the vesuvianite xenoblasts, are set in a fine granular mosaic of diopside, orthoclase, quartz, prehnite, and a little epidote. Patches of anisotropic garnet also occur, and it is possible that this mineral, as well as the prehnite and epidote, belong to the period of metasomatism. The vesuvianite shows extensive alteration into prehnite and a fibrous mineral described below (p. 42).
This assemblage occurs in bands which alternate with a prehnite-apophyllite assemblage, and it is believed (see p. 40) that prior to metasomatism these latter seams were represented by a wollastonite-orthoclase-diopside hornfels.
(xiv) Wollastonite-diopside Hornfelses.
Examples of this type are recorded from the granite contact near the head of Liddleton Creek, from Bonnie Blink Creek just above the junction of Liddleton Creek, from Moyne Creek, and from the diorite contact on the river below Deep Ravine. As most of these rocks have suffered some form of metasomatism, they will be considered only very briefly here. They find a place in this section, however, for it is evident that many of the metasomatized hornfelses were originally of this type.
In the hand-specimen they are fine-grained, greenish, granular rocks very rich in quartz. Wollastonite varies in amount and sometimes may be so abundant as to make up most of the rock. When less plentiful it may occur in small radiating masses or in bands parallel to the original bedding. In the field it is evident that the brachiopod shells have been replaced by wollastonite which, on weathered surfaces, is now represented by a fibrous, white earthy mineral.
32 PETROLOGY OF THE HARTLEY DISTRICT. iii,
This mineral has not been identified. It is isotropic and has a very low refractive index (1:460). Under the microscope masses of radiating wollastonite measuring up to 5 mm. across are set in a fine (about 0-1 mm.) granoblastic groundmass of diopside, quartz, orthoclase, and sometimes a trace cf plagioclase. Sphene and iron-ores are usually present as accessories. Alteration of the wollastonite into the unknown fibrous mineral seems to occur only on weathered surfaces, but in the body of the rock it commonly shows alteration into carbonates. Small andradite veins often thread through these hornfelses, but the occurrence of this mineral is referred to below. Scapolite is sometimes associated, but this, too, is regarded as a product of metasomatism.
A partial analysis has been made of a rock from Bonnie Blink Creek. This hornfels contains traces of scapolite and plagioclase.
SiO, 73°16