Ae (18 ts ht ae oy anne 40 oe) pea a 07 ball 4)" rid ¥ iy We a Hy Mey MA Shige vlutah tetas es his) oe pe Bevin A aN ia if ae oe yeti ah Pane iy — : us / y SOM athiity th ope Mm Abeal vi i sia a on ae aoe Mand ; at i i My nies ae Cate aha acs aeean ia arate tae reat sas ictal tat ar ty i i Ef oe 8 Ke ae ty ae ils ee inet : i ne ut Peas ee Pile, Pa Heal ‘ pth r f : ¥ " ee, ¥ ra ts cnet renee sn tytn oe tt bid Ay ey sh ) Hes vs fh y + ot Ay iron Reet ae Me an pied ‘ Hed ie Bia i hi aoe Wenner 7 nt ee ia ate On. Tiegh vas es Pg A vite ee et i see! ee “sh Kit Hy Wot) Ba a aie vat Hey ens > : ’ Lanes — 1 cf 1S 1 a 5 A i ; " z ; 4 os : 7 my £ i ; ‘ 7 Fe amt) P y ; ; 5 om. f , i re i Cae IN, i ; Tet HR : : Le f s by, } S ; | 4 oh a (Ai r : x : ae “}d / i | r vy i . u Sate 7 ol : : J bo 5 1 and ie 7 —— 3 ee : : \ - ~ 5 + Te OS " : - \ + \ De \ : tore i _ Ss ‘ : Ly Set -_ ‘ a . ; = i U ford ; : ' ' id ‘ y rs z A is a7 Lissa 2) 3 ara) =) i 7 > : : : r we Wy 1 pen ' " M ‘ . im ; 4 P ee } fi - ‘J | - 4 . 7 % r 7 oy é - I 1 y 5 i Le a F : } e a, 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- PRESIDENTIAL ADDRESS. ix 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 x PRESIDENTIAL ADDRESS. 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. PRESIDENTIAL ADDRESS. xi 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. xii PRESIDENTIAL ADDRESS. 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 xiv PRESIDENTIAL ADDRESS. 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 PRESIDENTIAL ADDRESS. xv 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% Xvi PRESIDENTIAL ADDRESS. 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. PRESIDENTIAL ADDRESS. XVil 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 xviii PRESIDENTIAL ADDRESS. 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) PRESIDENTIAL ADDRESS. Ks 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.) XXiv PRESIDENTIAL ADDRESS. (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 PRESIDENTIAL ADDRESS. xXV 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 XXvi PRESIDENTIAL ADDRESS. 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. ,1912.—L’Eau de Mer, Milieu Organique. Paris. 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. ,1930.—Die Osmoregulation wasserlebender Tiere. Biol. Rev. Camb., 5, pp. 309-356. 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. , 1932.—Water Regulation and its Evolution in the Fishes. Quart. Rev. Biol., Us INOS 15 19s) ISAs SuMNeER, F. B., 1906.—The Physiological Effects upon Fishes of Changes in the density and Salinity of Water. Bull. U.S. Bur. Fish., .25, pp. 53-108. WILLIAMS, T., 1852.—On the Blood Proper and Chylaqueous Fluid of Invertebrate Animals. Phil. Trans. Roy. Soc., 142, pp. 595-653. 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. 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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- 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 ) (° C.) 43 hours as aid 1:07 1-79 43 hours Se as — 1:89 6 days .. oe sie 1:20 iLo'7/35 Thus the results for Australian species combine with many of those obtained by investigators elsewhere to show that a “law of poikilosmoticity” is by no means applicable to all marine invertebrates when the surrounding water is of lower salinity than ordinary sea-water. It is, however, of greater validity when the external concentration is increased above that of ocean sea-water. Both Frédéricq and Duval found isotonicity in different species of crabs under conditions of increased salinity. The former found identity of internal and external freezing points for Carcinus maenas when immersed for three days in water of A 3:11° and A 3:84°. The latter obtained his result from Platycarcinus BY ENID EDMONDS. 245 pagurus, Palinurus vulgaris, Maja squinado and Carcinus maenas immersed in various solutions which froze between —2° and -3°. The crab Heloecius cordiformis was examined under conditions of increased external concentration. It can endure a large increase in the concentration of the external water, and will live for some time at least in double strength sea-water. The previous notation for strength of solution will be used for setting out these results. Thus a 100% solution is one of normal concentration, 150% is a solution half of ordinary sea-water and half of double strength sea-water, and so on. When Heloecius was first put into concentrated sea-water the specimens were left only for a day or two before the freezing point of the blood was determined, as this time was all that was necessary for the completion of the changes in dilute solutions. The results were very surprising in comparison with those of Duval, which showed isotonicity for Carcinus in 26 hours. In water of nearly double strength (A 4:14°) the blood of Heloecius gave a A equal only to 2:50°. Again, for a 150% solution of A 3-:14°, the A for the blood was 2:18° on one occasion of immersion for 88 hours, and 2:29° on another when the experiment lasted 42 hours. A large number of experiments were subsequently carried out, and they all showed that the blood remains hypotonic to a large degree when the external medium is concentrated above the normal, and the time of immersion only one or two days. It was now necessary to determine whether this hypotonicity was permanent, or whether it was due to a slower rate of change than under conditions of dilution. The latter alternative was found to be the true one. Examination over a long period showed that the internal change is at first fast, but becomes gradually slower. Instead of ceasing altogether after a day or so (as is the case for dilutions) the change continues at a slow rate, until the salinities of the inner and outer media are almost equal (the difference being no larger than that for crabs in ordinary sea-water). The time necessary for the completion of the change was not constant, but varied from two to five weeks. The rate of change can best be seen from the following tables and the graph (Fig. 5), which unite the results of typical experiments, TABLE 11. Heloecius cordiformis Immersed in Solutions of A =3-30-3-40. Difference between Duration of A of Blood. the Two Media. Experiment. € C.) (° C.) 0 ae 1:83 1:49 4 hours 1-98 1-34 21 hours 2-49 0-89 28 hours 2-19 1-21 5 days .. 2:54 0-78 6 days .. 2-62 0-70 14 days .. 2-56 0-82 21 days .. PACs 0-68 21 days .. 2-85 0-55 48 days .. 3:41 & * The external concentration would have been a little altered by evaporation and, as it was not obtained at the end of the experiment this difference must be omitted. 246 RELATIONS BETWEEN INTERNAL FLUID AND WATER OF ENVIRONMENT, TABLE 12. Heloecius cordiformis Immersed in Various Solutions of Increased Concentration. Duration of A of Solution. A of Blood. Experiment. (2p) (E04) 6 days .. Po By7/ 1°96 7 days .. 2-64 2:18 28 days .. 3°28 2:92 29 days .. 3:17 3°15 36 days .. 3:24 3:10 36 days .. 2-42 2-28 o a uw % i FREEZING POINT ‘OF BLOOD ow bs + o ° e 4 6 t2 e+ 43 DURATION OF EXPERIMENT (CIN DAYS) Text-fig. 5.—Heloecius cordiformis immersed in solutions of A 3:30-3-40. Thus we find that H. cordiformis agrees with the marine invertebrates previously investigated in that, after an increase in the concentration of the surrounding water, its blood comes into the same relation with that water as it was originally with the ordinary sea-water. It differs considerably, however, from the crabs examined by Duval and Frédéricq in the length of the time taken to complete the internal change. SUMMARY. A number of species of Australian crabs from diverse habitats, such as a typical ocean coast, estuarine flats, and fresh waters, have been examined in order to determine the relationship of their body fluids to the external medium under natural and under experimental conditions. Five species of crabs have been found to be homoiosmotic in diluted sea-water. This gives much additional support to the view that the condition may be regarded as general for the crustacea. The number of species of marine crabs in which, under normal circumstances, there is a distinct anisotonicity between the body fluids and the external medium, has been increased. The difference is on the side of hypotonicity. BY ENID EDMONDS. 247 It is noticeable that, as in other cases, Heloecius cordiformis survives in highly diluted sea-water for a much longer time than in fresh water. The effect of the trace of salts is important. The osmotic pressure of the blood varies amongst the individuals of any one species when taken from the same conditions and the same locality. Although Heloecius cordiformis behaves like certain previously examined European crustacea in that its blood comes ultimately to a poikilosmotic condition when the crab is placed in water of increased concentration above normal ocean salinity, it differs from the known cases in taking about a month to reach this condition in highly concentrated solutions. Bibliography. BAUMBERGER, J. P., and OtMstTEDT, J. M. D., 1928.—Changes in the Osmotic Pressure and Water Content of Crabs during Molt Cycle. Physiol. Zool., 1, p. 531. Borrazzi, F., 1897.—La Pression Osmotique du Sang des Animaux Marins. Arch. Ital. Biol., 28, pp. 61-72. DAKIN, W. J., 1908.—Variations in the Osmotic Concentration of the Blood caused by Changes in the External Medium. Bio-Chem. Journ., 3, pp. 473-490. , 1912.—Aquatic Animals and Their Environment. Internt. Rev. ges. Hydrobiol., 4, pp. 53-80. ,and EpMoNDs, ENID, 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. Exp. Biol., 8, pp. 169-187. Duvau, M., 1925.—Recherches physico-chemiques et physiol. 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. FREDERICQ, L., 1901.—Sur la perméabilité de la membrane branchiale. Bull. Acad. Roy. Belg., Sér. 3, Tome 30. —, 1904.—Sur la concentration moléculaire du sang et des tissus des animaux aquatiques. Archiv. Biol., 20, pp. 708-730. Mont, R., 1914.—La variabilita della pressione osmotica nelle diverse specie animali. Jie SOG, TEC ISOS INChies, Be 1s Beale SCHLIEPER, C., 1930.—Die Osmoregulation wasserlebender Tiere. Biol. Rev. Camb., 5, pp. 309-356. MISCELLANEOUS NOTES ON AUSTRALIAN DIPTERA. III. By G. H. Harpy, Queensland University, Brisbane. [Read 31st July, 1935.] Subfamily CuRYSOSOMATINAE. In my catalogue of the Dolichopodidae, I drew attention to the need for improvement in the treatment of genus Chrysosoma and allies. I gave in the key the treatment usually adopted, but did not follow the system when arranging species under the genera. I was unaware of the paper by M. l’Abbé O. Parent redescribing two of Macquart’s types, for the periodical containing that paper is not in any Australian library. A second paper by this author came to hand when the catalogue was going through the press and I was able to refer to new species described there. Much has been done since by Parent, and I append a list of the papers that contain references to Australian species as far as I know them. Half of these papers are not accessible in Australian libraries at the present time. It is necessary here to point out that Parent uses Becker’s system of classification, and hence he and I place species in quite different genera. He has found that the antennal structure is ambiguous in a minute percentage of specimens, whereas, dealing only with the Australian material, I find this ambiguity in a big percentage. I therefore divide the species into natural groups which are defined as far as possible. Key to genera of the Chrysosomatinae. 1. Frons deeply excavated between eyes. Wings usually with second median vein strongly indicated (i.e., fourth vein forked), but may be missing ........... Blake O-cnorc OI Gsa DES OPLOREYSrReTa ROTO lon Orono OeROT a RGR Een Ero ba Gerona o ns see mG CHRYSOSOMATINAE. 2 Frons slightly or not excavated. Second median vein absent. Rarely do these characters occur, then if frons be excavated the radial veins all end at costa well before the apex of the wing, and if the second median vein is indicated there is also an appendix at the bend of the first median. In both cases the hind tibiae have many strong bristles which are about as long as the thickness OF FUE ETO Laos aie a ep ecceicns vattonresseus ee me dane oes ete deat TS GAS ciIe te ORS ES SONI TEE eh oe Other subfamilies. 2. Second median vein entirely eliminated and the first median gently curved ...... 3 Second median vein present usually and the first median branches away abruptly .. 4 oe eAbdomen Short, swinssenonmMalamn arial cieicicicr icteric ioe eicesae Mesorhaga Schin. Abdomen jlones wines weLry, NALLOWs! ey ccee otic coe ceinciseeicieinns Australiola Par. 4. First median vein strongly sinuous at its basal half. Antennae with a swelling on basal segment forming a long process .................- Megistostylus Bigot. First median vein only bowed or straight. Antennae without a process at basal FSh=}=9) 00XS) Ch ea Bn rath er OREM GR erO COTA D Glan Den SIC CRON COR A RC IPSEIa Tria GiaiAicro cd'o'o oo 5 5. Male with the first radial vein very long, reaching costa at a point beyond that above the apex of the median cell. Male with hook-shaped cilia on costa..... ats vs emepatlamants do ake. fav's nay ray euawee Keeley omen ARON Wo ne te Roper ob sree tela a Sea renee Ee eecPe Parentia, n. gen. Male with the first radial vein short and the costa not ciliated ................ 6 6. First median vein strongly bent to a right angle. Median cross-vein strongly sinuous and often with a veinlet in centre or somewhat angulated there ....... CGN CUCROTO 1-5, 0) OCOLDICHCPU NG CHER VOL Eh OLOTO cha OANIG Oto ExE GA ATO TC ERO GER TES BIE OO OIA Heteropsilopus Big. BY G. H. HARDY. 249 7. Antennae with a long conical third segment and a terminal arista. A well developed sinuous median cross-vein and two pairs of scutellar bristles are usually DGES CT arnwegehonceon Rome eh ei anortonercel inch eaeirs musysaetewahe. eleanor aioe R arn Pehoiee Chrysosoma Gueér. Antennae normally short and with a dorsally placed arista but variable. It may reach the length of one and a half times or even twice as long as thick with a terminal arista. Other characters variable ................. Sciapus Zell. ScIaAPUS complex. It seems necessary to review the position of this complex as far as it affects the Australian fauna, the names and synonyms being as follows: Sciapus Zell. 1842 (Sciopus of authors) with type platypterus Fab., Hurope, includes Leptops Fall. 1823 (preoccupied), Psilopuws Meig. 1824, Psilopodinus Bigot 1840, and Psilopodius Rond. 1861. Chrysosoma Guérin 1832, and Agnosoma Guérin 1838, type maculipennis Guérin, from New Guinea, would seem to have as synonyms Oariostylus Bigot 1859, Mesoblepharus Bigot 1859, Tylochaetus Bigot 1888, Spathipsilopus Bigot 1890. Oariopherus Bigot 1890 and Hudasypus Bigot 1890. Heteropsilopus Bigot 1858 can be isolated as a definite concept with type grandis Macq., and possibly Plagiozopelina Engel 1912 as a synonym. The genus Condylostylus Bigot 1859, type bitwberculatus Macq. from Brazil, forms a good concept that seems to have little in common with the Australian material and so Australian forms placed under it revert to Sciapus. There are a number of other generic names proposed but founded on American forms that do not seem related intimately with those of Australia and so are ignored here. Nevertheless, I can detect six main groups in the Indian and Australian forms that seem to warrant names in accordance with the following key: 1. First radial vein reaching to and beyond a point level with apex of median cell Onemthe malerae weasel! Mays hee LN ee URE ERE TR. ey ee at Adah RE Ren aur red 2 MMEStnaGlaluavietme GShioritey ell ahews Mecenscd lors a: ve.m ens aaj violet el evel aire paired hoped aa ECC aes 3 DB Come, Giles! Win wR Sdsoacocubo0g Gist=sroup) ene eee Parentia, n. gen. Cosi: Tao GWEC! odocaoncocoounogcd (AinGl EAeOWO)) Gaanacooc Type, liber Par., Fiji. 8. Costa ciliated, wings slender ...... (38rd group) .. Type, adhaerens Beck., India. Costasnotnciliateda OrmranrelypiSOmes concrete & shsctnsplmopaisie: cuciseecuemeko ee Coa eis ot ssiebew se say ue ee ee 4 4. Median cross-vein strongly sinuous and more or less angulated in centre, often with a veinlet there. First median vein bowed to a right angle .............. PF Ne ay Pe SO Retro raked? jayne nickels ara Ne (4th group) ......... Heteropsilopus Big. IWaAthOutmtheseucharactersm combined) aasccs ee acetone eae ene neice D 5. Third segment of antennae long and strongly conical with an apical arista. Median CHORE SinwiOws) Al IEGAGE WIKWANhA SO SNcacldcldccccoscdblauoacavaubadoussocboseas Ha aaa Biot tin meee oh Sek ato ca nope ue mabe Oia (th sroup)” 4e.2.59.5. .ChnvsosomaGuéer: Third segment of antennae variable, usually short, arista apical or dorsal. Median GQORSAVENN OE GimoOwMs lowe Wiebke Gioeblue “soos cgcboopcaccocnsoubeooanuaun SHOES ON EC, BROUCEU OS SOREEOAID CEERI acne RO Cua (6th sroup)) Fyre ees ClapncSmZelle The 5th and 6th groups are heterogeneous and I think Mesoblepharus Bigot, type senegalensis Macq. and synonym Hudasypus Big., might make a nucleus for another group, possibly incorporating the Australian interruptum Beck. PARENTIA, nN. gen. Type, Condylostylus separatus Parent. Tasmania. The arista is placed dorsally or apically on a short or rather short third antennal segment. Normally the scutellum has two pairs of bristles. The wings have the first radial vein on the male unusually long, reaching beyond the level of the apex of the median cell and in addition there is a fringe of rather long hook-shaped cilia along the costa (illustrated by Becker, 1922, fig. 203). The female has a short radial vein and is without the cilia. The forms are all dark Kk 250 NOLES ON AUSTRALIAN DIPTERA. iii, blue-green, except the typical form which seems to have colour dimorphism in this respect. This genus is well represented in New Guinea and may occur beyond that area. It is not known from New Zealand and India, where another group with ciliated costa seems to take its place. Key to males of species of Parentia. 1. Wings with a duplicated row of cilia on costa ................ duplociliata Par. WinestwithwawsineleurowsofaciliamonecOStan mE Rinne enone cena 2 2. With wings dark and hairs abnormally abundant. Legs entirely dark ......... SORT EL ROR HL ELCRERODY CUBANA KROME art Ra Crate on Ga Cae at ac SCE aN nigropilosa Macq. Winssehyalines Normally ehairedsispeciesmts ies ieloela Rta elena eee 3 oo Peeks entirely: ark eee sic seeyeteyeus seeeesuene Gy conscious ia fetemeds den duehipeewe) Ghelesemsieonee ete Meter semen eas 4 hers partly lsht Coloured) — .acecihvecscs sida cotene evens ooeualig thieves Cl av nisus th entiers) E) nathan 5 4. Anterior femora with long black bristles on ventral surface ....... tricolor Walk. Anterior femora with only yellow or white hairs on ventral surface .... dubia Par. 5. Femora rather widely yellow-brown at apex ................20+.... separata Par. Remora entirely dark, except perhaps at the tip ................ dispar Macq. PARENTIA DUPLOCILIATA (Parent). Chrysosoma duwplociliatum Parent, Ann. Soc. Sci. Brucelles, (B) liii, 1933, 172. Hab.—Northern Territory. PARENTIA NIGROPILOSA (Macquart). Psilopus nigropilosus Macquart, Dipt. Haot., suppl. 2, 1847, 56.—Sciapus nigropilosus White, Proc. Roy. Soc. Tasmania, 1916, 251.—Condylostylus nigro- pilosus Hardy, Aust. Zool., vi, 1930, 131; Parent, Ann. Soc. Sci. Bruxelles, (B) 1ii, 1932, 126. Determination of this species is based on White’s identification, but there is no assurance that White identified the species correctly. This list of references may cover a complex. Only the male is known to me. Hab.—Tasmania. PARENTIA TRICOLOR (Walker). Psilopus tricolor Walker, Ent. Mag., ii, 1835, 471—Psilopus gemmans Walker, List Dipt. B. Mus., lii, 1849, 644; Parent, Ann. Mag. Nat. Hist. (10), xiii, 1934, 34; and Ann. Soc. Sci. Bruxelles, (B) liii, 1933, 178—Condylostylus amoenus Becker, Cap. Zool., i, 1922, 219, fig. 203; Hardy, Awst. Zool., vi, 1930, 131; Parent, Ann. Soc. Sci. Brucwelles, (B) lii, 1932, 126. Walker’s description fits well this common species, so I am giving preference to the name tricolor. Parent found that the type of gemmans was conspecific with Becker’s species. Both sexes are before me. Hab.—New South Wales and Victoria. Walker and Parent also record it from Western Australia. PARENTIA DUBIA (Parent). Chrysosoma dubium Parent, Ann. Soc. Sci. Bruxelles, (B) xlix, 1929, 201, figs. 50, 51; and lii, 1932, 109—Condylostylus dubius Parent, Ann. Soc. Sci: Bruzetles, (B) lii, 1932; 126. A Queensland species before me agrees fairly well with the description of this one and runs to it in the key, but differs in having the lamellae very long and bifid. Both sexes are before me. I have not seen Parent’s form. Hab.—South Australia. PARENTIA SEPARATA Parent. Condylostylus sp., Hardy, Aust. Zool., vi, 1930, 130 (in key) —Condylostylus separatus Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 127, fig. 19. BY G. H. HARDY. 251 The only females I have been able to associate with this species have the femora and tibiae entirely yellow. These occur together and are quite common; I have not seen females with legs like those of the male or males with legs like those of the females. Also the female is green in colour. Hab.—Tasmania: Generally distributed over the eastern half of the island from December to March. Victoria: Common in the Melbourne district. PARENTIA DISPAR Macquart. Psilopus dispar Macquart, Dipt. Hwot., suppl. 4, 1849, 125.—Sciapus dispar White, Proc. Roy. Soc. Tasmania, 1916, 251—Chrysosoma dispar Parent, Ann. Soc. Sci. Bruxelles (Vol. Jub.), xlvi, 1926, 18; and (B) lii, 1932, 109. Hab.—New South Wales. CHRySsosoMA Guérin. Chrysosoma Guérin, Voy. Coq. Zool., 1831, Atlas, Tab. xx, 25, vii. The species I place in this genus have the third segment of the antennae at least one and a half times longer than broad and the very long conical appear- ance with the arista placed terminally. The median cross-vein is sinuous on all described forms and probably all species with the straight median cross-vein are best relegated to Sciapus until its true associations can be worked out. Doubtless Chrysosoma as here understood is a complex group; nevertheless there seems to be a general alliance between the majority of them. Key to species of Chrysosoma, based mainly on males. PATI StaywithwanspauuUlate rai excgin agsuctcbe rolsicsl aisle relior als ojpa a Seven edeeken suse callosum Parent. Arista simple, at most slightly flattened at apex and then white in that area ... 2 2. Wings entirely fuscous or almost so. Long black hairs on frons .. funerale Parent. Wings with fuscous markings. Costa ciliated .............. interruptum Becker. AVVenl' eS Salar DIT OF Moe pacso st sere eer vee cee a) tecteod en 7satsangoen BGiai Gulet al lauds touts SyeMeanetie) eatemettol: italic Rapawaleyre peers Rome h opens 3 3. Intermediate legs on male with the fourth tarsal segment slightly enlarged and longer than the two prior segments united .............. . caelicum Parent. intermediatesless! of male) withitarsi not So wLormed. Sse sence) iether terete: 4 4. Anterior femora yellow, the others black .................... diversicolor Parent. All femora yellow or practically so. Hind tibiae with a black ring on male. Fourth segment of intermediate tarsi white on male ......... leucopogon Wiedemann. HETEROPSILOPUS Bigot. Bigot, Ann. Soc. Hnt. France, (3) iii, 1859, pp. 215, 224. Type, by original designation, Psilopus grandis Macq., Australia. A natural group is formed by cingulipes (syn. grandis) and associated species distinguishable by characters given in the key to genera. Key to species of Heteropsilopus. 1. Wings clear. Arista subapical on a very short third segment. Two pairs of Soule LIMOS “oooseeoaneanbsedonasouadansnocabodacvics cingulipes Walker. Wings marked. Usually a dorsal arista and one pair of scutellar bristles ...... 2 Me Wwohovas Iieinohy Greeley) eylomee Viens “Secocasabosnouddvoacagndots jacquelinei Parent. Wings with spots limited to cross-veins and any shading elsewhere exceedingly SPER UTA Chminerevenene liens csncnsue ee atetemer atiectarion eve wtraliaoe Dice cvialiled alte lence ae tude Qeveaganten alienate brevicornis Macq. Wines swithywelltnanrkwe GmEAasciay , lhe sires eee ein nv cetera jean) semua oie Lennar gros vase et eure tetioneie 3 3. Apical segments of intermediate tarsi peculiarly formed on male. With conspicuous DLIStIESPOMPULI DIA Machin iavecetan exe oy ices joes ialisis es ste eyfeitel se siren eviovcmeney ees ingenuus Brichson. Segments of intermediate tarsi more normal but with a fringe of cilia. With rather INCONSPICUOUSPHLISLIESHONMELDIAeMy wer. /-saeieiet-l eileen stn tease plumifer Beck. HETEROPSILOPUS CINGULIPES Walker. Psilopus cingulipes Walker, Ent. Mag., ii, 1835, 472; Parent, Ann. Mag. Nat. Hist., (10) xiii, 1934, 9—Chrysosoma cingulipes Hardy, Aust. Zool., vi, 1930, 126.— Psilopus sydneyensis Macquart, Dipt. Exot., suppl. 1, 1846, P, xi, f. 16.—Psilopus 252 NOTES ON AUSTRALIAN DIPTERA. iii, sidneyensis Macquart, ibid., suppl. 2, 1847, 56; White, Proc. Roy. Soc. Tasmania, 1916, 251.—Psilopus grandis Macquart, ibid., suppl. 4, 1849, 126; Parent, Ann. Soc. Sci. Brucelles, (B) lii, 1932, 231 (synonymy).—Psilopus eximius Walker, Ins. Saund. Dipt., i, 1852, 209; Parent, Ann. Mag. Nat. Hist., (10) xiii, 1934, 16.— Psilopus angulosus Bigot, Ann. Soc. Ent. France, (6) x, 1890, 285; Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 216—Chrysosoma alatum Becker, Cap. Zool., i, (4), 1922, 188, fig. 159; Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 109.— Chrysosoma micans Parent, ibid., 1932, 109..—? Chrysosoma metallicum Parent, ibid., 1932, 113. Much of the above synonymy is recognized by Parent who added sydneyensis Macq. and micans Par. to the list. The new synonymy is angulosus Big. and metallicum Par. I have a specimen of the latter, but regard it as a variation and it will require a male before it can be established definitely as a distinct species; meanwhile it seems to me advisable to place the name as a possible synonym. Hab.—Queensland to Victoria. HETEROPSILOPUS BREVICORNIS Macquart. Psilopus brevicornis Macq., Dipt. Exot., suppl. 4, 1849, 124—Sciapus brevicornis White, Proc. Roy. Soc. Tasmania, 1916, 249; Hardy, Awst. Zool., vi, 1930, 126; Parent, Ann. Soc. Sci. Bruxelles (Vol. Jub.), 1926, 16; and (B) lii, 1932, 117.—? Psilopus venustus Walker, Ins. Saund. Dipt., i, 1858, 209; Parent, Ann. Mag. Nat. Hist., (10) xiii, 1934, 36—Psilopus chrysurgus Schiner, Novara Reise Dipt., 18638, 214—Chrysosoma chrysurgum Becker, Cap. Zool., i, (4), 1922, 172; Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 109; Hardy, Aust. Zool., vi, 1930, 126.—Sciapus chalceus White, Proc. Roy. Soc. Tasmania, 1916, 250.—Chrysosoma volucre Becker, Cap. Zool., i, (4), 1922, -142, figs. 74-6; Hardy, Awst. Zool., vi, 1930, 126.—Sciopus bimaculatus Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 117, figs. 7-9. The above synonymy is new. Parent agrees with me, in a letter, that his form is the same as Becker’s, but is not yet prepared to give assurance that these are identical with Macquart’s type which is incomplete. Nevertheless, he writes that he can find nothing to disagree with this synonymy in the descriptions. The names given by Walker, Schiner, and White, according to the descriptions, would also fall to synonymy, and there can be no doubt in this respect concerning Schiner’s description, whilst that of White applies evidently to a variation. In describing venustus, Walker gives the characters of a male with wing marks, whereas Parent, redescribing from Walker’s material, refers to a female without wing marks, missing the appendix to the median cross-vein, but apparently agreeing in other respects. Hab.—New South Wales to Tasmania. Records would indicate that this species occurs widely over Australia. HETEROPSILOPUS INGENUUS Hrichson. Psilopus ingenuus Erichson, Arch. f. Nat., xiii, 1842, 273—Sciapus ingenius Hardy, Aust. Zool., vi, 1930, 127—Sciapus trifasciatus White, Proc. Roy. Soc. Tasmania, 1916, 248; nec Macquart, 1849.—Sciopus gloriosus Parent, Ann. Soc. Sci. Bruwetles, lil, 19382, 119: The above synonymy is amended from that of my catalogue, with Parent’s name added as a new synonym. Hab.—Tasmania (abundant) and Victoria. The species is plentiful in the Melbourne district, and there are females before me from Adelaide, South BY G. H. HARDY. 253 Australia, and from the extreme north (Tooloom) of New South Wales, and these apparently are the same species. HETEROPSILOPUS TRIFASCIATUS Macquart. Psilopus trifasciatus Macquart, Dipt. Exot., suppl. 4, 1849, 126—Chrysosoma trifasciatum Becker, Cap. Zool., i, (4), 1922, 176; Parent, Ann. Soc. Sci. Brucelles, (B) Iii, 1932, 109—Sciopus trifasciatus Parent, Bull. Mus. Hist. Nat., (2) iv, 1932, 879; and Ann. Soc. Sci. Bruxelles, (B) -liii, 1933, 179. By comparison of the figures with specimens of ingenuus, I conclude that Parent has been misled in regarding trifasciatus Macq. as distinct from ingenuus. It seems probable that his figure, made from one of Macquart’s specimens, is the result of faulty interpretation due to the specimen being in poor condition and not to differences in actual structure. This matter needs elucidating, but in the meanwhile the above references are kept separate. HETEROPSILOPUS PLUMIFER Becker. Sciapus plumifer Becker, Cap. Zool., i, (4), 1922, 206, figs. 183-4; Parent, Ann. Soc. Sci. Brucetles, (B)) iii, 1932) 122. From near Becker’s type locality comes a form that is to be distinguished from ingenuus Er. by structures, some of which are mentioned in Becker’s description, and this form doubtless will prove conspecific with plumifer. The anterior tarsi are much longer than those on Erichson’s species, being one and a half times longer than the anterior tibiae. This occurs on both sexes, and the male has the posterior tarsi similarly much longer. Also the tibiae are relatively bristleless in appearance, the bristles being small and yellow instead of well developed and black. In addition the male has the intermediate tarsi ciliated for their complete length and none of the segments are otherwise ornamented. Hab.—New South Wales: Blue Mts. Sciapus Zeller. Zeller, Isis, xi, 1842, 831. Other than those with wing markings, there are very few species below that lend themselves to ready recognition, but I have made an attempt to give a key that will aid in the determination of species. Many forms are known only from the female, and the species I have been able to identify are marked with an asterisk (*). IKkey to species of Sciapus. 1. Wings with distinct markings. Median cross-vein straight .................... 2 Wings without markings, clear or more or less suffused with grey ............. 5 A, IMO LRH qOlaecl ie Was lI GNOME CORMA, touconadovouadcgonnoansopon nanos o00 3 IN(OTES Owe TI) ATs OCD aes cis alts remtiae Nowra eases sy ticmaenar le Susimyicesasy dene Dea anes Sires se ep ab ctice cima ew nano Ten area tn epioare fret: 4 3. Third radial vein distinctly though slightly sinuous. Hypopygium long, with laterally directed and rounded lamellae .................. *connexus Walk. Third radial vein not sinuous. Hypopygium short and lamellae apically directed AMNAGDOIMUCCAS ray casesieots usd etc as, Sy aprssians ale eee (6a dayrauiside stages aieeee etn I oReRE Reese proximus Par. 4. Wings with two fascia across wings, usually complete and basal one not quite reaching costa. Lamellae not exserted beyond apex of hypopygium .......... NN Se ATIR PRS RE ay Sey eau SEL OA Sy fod RN spun ae Salleoas ee Adaliasatiene sted tele meee *discretifasciatus Macq. Wings with interrupted bands, making four spots, two of which touch costa. Lamellae long, exserted well beyond apex of hypopygium ........ quadrimaculatus Par. HSV an CV OSS=VELMSIMUOUSiis kopeere ves ecre lactis oe erie sap Creep sua caee es ona ete esc Lee CR eaN UA MTT Ae ereeue tauren 6 Medianmcross-veln straishnt wor practically, Som nicer cicne sienna eienicne etanaiens 8 GS, PAU CORAS RE DTACKS Reece ccs cavers ee ease. eve ha im sat ta eal al racic os ASE EE ORIG EATERS) Sete tA 8 ne NE 7 ANIECELON CORA CVC lL O Way carat across eter cta ce seeetr sh es STEN Sy eT OG CRA ICU es CS lee difficilis Par. ake SSXOTREW covey; OMEN) Fer iclaren en stens iia os tohcmeaeesbes Hath ats ernea cr otnestorone crolbror so Sin talc ata osetia es imparile Par. Squama yellow. Male with a fringe of cilia on intermediate tarsi ..... nobile Par. 254 NOTES ON AUSTRALIAN DIPTERA. iii, 8. Legs entirely black. Base of fifth radial vein recurrent ...... *qustralensis Schin. Legs otherwise coloured. Radial vein not recurrent at base ..........-.-+--:+-- 9 9. KFemora black or metallic-green, but yellow at apex ....... *nigrofasciatus Macq. Nera, Gmummeahy Wallon: casosodavoccssuSdodoOadoub go dco noo oe oO UNDE NO OBOAG OAD 10 10. Hypopygium with elongated processes on lamellae ............-+2ee esses eeee Tal Hypopygium presumably without such processes .......----+..---+-:eeesese sees 12 thal, NANO TANKS ORWIRS) @ie WIN JOHOCESSES saocaaccoconceccsnaaccgecbood anonalicornis Beck. IWithethreespalnsnoLsulChmproGeSSeSmmrestn let ieici-len Rleisiecneleiieneter *triscutatus Hardy. 12. Third radial vein strongly bent downwards towards the first median vein and away from the second radial. First median vein branches remote from the median cross-vein by about one and a half times the length of the latter .......... Ee RO eH SOLS Ore RCRA EE oe ch carOee OF TRIES Igwy. Ge cee ty SPokerC utonors io asace graciliventris Par. Wii NOMEN Vaouiaddooosa dey oodob moos oH audd A doU Oe oa DDO OwWln.aO DOC ODOC O90 3 1/98 “Anitennaeventirelys Place fee cake as eels cel ele e leue aneteusls tale due Lemon ellswel tenets siete elle lees tele ner memes 14 Antennaeysvellow: (PELE sibel eielone cildiedsp sta eyetae obs eiceautstekekchancuelletedsiciakeitan te teuc tlic e at tanete 16 HEY: Cen 2GraWeYatsn loves (oleae MA IMR Cra RM iC Woe a Pitre cota G Gi tee eo crete cr OG ClO CD One Gado Be sublectus Walk. INE WHOSE Oalhy VINA Oe ORUeCIOM Ike IG soonncnsabcocaob udp eso ME bO DSO aOD 15 Le MeMAcenparallel—Sid Git yes, sate one ssceene ln eosrese le erode sues) = siteiley steel. etellaetie ne ite chetiscutatus Par. TDR COMPETI TYCO sonasoaasooudnouggsonoogDoooapUaoD SoS nigrociliatus Par. 16. Hypopygium with four long sinuous apical bristles on each lamella .. *sordidws Par. JEHY aon Sivoo joyrresyonaneyalhyy TaeIE Yo) THONMONS! 5 Bs agocnsbuedoadbondoOO Odo ODE hoo D OOD ALY 17. First median vein bent, with a low circular arch, practically a quadrant ....... OEE a SE On CRT oe CCR ONCE EER EL Pace ER CHE Oras COMIC amc CeO ORC oS C BDROROT AICO O On iD zonatus Par. Hirst median vein bent to a broad rectangular arch ~.....:.-....-.. *mollis Par. SCIAPUS NIGROFASCIATUS Macquart. Psilopus nigrofasciatus Macquart, Dipt. Hzot., suppl. 4, 1849, 126.— Condylostylus nigrofasciatus Parent, Bull. Mus. Nat. Hist. Paris, (2), iv, 1932, 876.—Psilopus viduus Schiner, Novara Reise Dipt., 1868, 216.—Condylostylus viduus Becker, Cap. Zool., i, (4), 1922, 220; Hardy, Aust. Zool., vi, 1930, 131; Parent, Ann. Soc. Sci. Brusvelles, lii, 1932, 127.—Chrysosoma regale Parent, ibid., Ibhis ale eyz alalals Schiner’s name is placed here as a new synonym. Both sexes are before me and the male agrees with the description of regale which was placed by Parent as a synonym of nigrofasciatus, after seeing the type. Hab.—New South Wales. SCIAPUS SUBLECTUS Walker. Psilopus sublectus Walker, Ins. Saund. Dipt., i, 1852, 211.—Condylostylus sublectus Parent, Ann. Mag. Nat. Hist., (10), xiii, 1934, 31. The identity of this species is unknown, and for its probable position in the key I depend entirely upon Walker’s description. Hab.— Tasmania. SCIAPUS SORDIDUS Parent. Sciopus sordidus Parent, Mitt. Zool. Mus. Hamburg, xliii, 1928, 193; Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 123; Hardy, Awst. Zool., vi, 1930, 132.— Sciapus anomalipennis Hardy, ibid., vi, 1930, 128, figs. 1, 2; Parent, Ann. Soc. Sci. Brucelles, (B) lii, 1932, 117. A male from Victoria agrees with S. anomalipennis Hardy, having identical characters except that the hypopygium is apparently larger and reaches the apex of the fourth abdominal segment, whereas on Queensland specimens it reaches to between the apex and middle of the fifth segment. Parent, who has only seen the female of his form and both sexes of mine, is in agreement with me, regarding them as conspecific. Hab.—Queensland to Victoria. A male from Carrum, in the latter State, is in the collection of Mr. F. EK. Wilson. BY G. H. HARDY. 255 HyYDROPIIORINAE. Already five genera recognized as occurring in Australia are listed under this subfamily, namely: Hydrophorus Fallen, Paraliptus Bezzi, Liparomyia White, Scorpiurus Parent, and Paranthinophilus Parent. To these must be added Thinophilus Wahlbg., recorded here for the first time. The genus was discovered by Mr. L. Wassell and myself when making an unsuccessful attempt to secure Paraliptus, only two specimens of which are hitherto known, both taken by Mr. Wassell at a light when on camping trips with a motor-boat, and both specimens were sent to the late Dr. E. W. Ferguson. HyYDROPHORUS PRAECOX Lehm. Parent, Ann. Soc. Sci. Bruxelles, (B) lii, 1932, 71. Records of this species are given by Parent from Canberra and New South Wales. Specimens from Sydney and Hobart are before me and were mentioned (erroneously as two species) in my catalogue without specific determination. The species conforms well with Lundbeck’s description (Diptera Danica, iv, 1912, p. 346). THINOPHILUS WASSELLI, Nn. Sp. do. The whole body is covered with a blue-green iridescence with purple tinges more or less obscured by a pulverulent olive-yellow. The antennae have the two basal segments yellow, the third black or mainly so, and the palpi also yellow. One pair each of vertical, ocellar and postvertical bristles all black, and one row of white postoculars that gives place to black towards the vertex and where extra bristles tend to form two rows, the second row numbering up to three bristles or may be absent. Some yellow and white hairs form a small scanty beard. Each side of the prothorax are four short black bristles placed in a row. The mesonotum is without hairs, except for a small group of short stiff ones that run into eight dorsocentral bristles, the last two only being strongly developed. Outside these there is a line of four bristly hairs reaching the trans- verse suture, beyond which, in the same line, two supra-alar bristles occur. One each of humeral, posthumeral, notopleural and postalar bristles stand isolated except for two bristly hairs on the humeral tubercle. Two pairs of bristles occur on the scutellum. Some scanty long hairs occur on the propleura anterior in position to the spiracle, otherwise the pleura is bare. The abdomen contains six normal large segments uniformly covered with black stiff hairs, followed by a complex of much reduced segments and the hypopygium which is mainly retracted into a groove on the venter and reflexed, but showing a Y-shape induced by two diverging slender parts reaching the fourth segment. The anterior coxae are yellow, with long black hairs placed anteriorly, and covered with a pulverulent white. The remainder of the anterior legs are similarly yellow except the apical tarsi, which are stained with black, the whole being covered with short scanty black hairs and only four bristles occur all on the anterior side of the tibiae. The intermediate and posterior coxae are yellow with a pulverulent grey that makes them unicolorous with the pleura, the remainder being coloured as the anterior legs, but the bristles of the tibiae are more plentiful and more generally distributed. The posterior coxae have a lateral bristle. The venation is typical. The female is similar, but only five abdominal segments are to be detected and on the anterior coxae only short black hairs occur. 256 NOTES ON AUSTRALIAN DIPTERA. iii. ” Hab.—Queensland: Southport, December, 1932, and January, 1933; 7 males and 12 females, occurring plentifully on the uncovered tidal mud around Mangrove swamps. Note—In Parent’s key, this species runs to Parathinophilus, but may be separated by the absence of acrostichal bristles and other characters. References. Harpy, G. H.—Australian Zoologist, vi, 1930, pp. 124-134. (Contains full references to literature except those of Parent’s papers.) PARENT, O.—Ann. Soc. Sci. Bruxelles, (B) xlvi, 1926, pp. 205-229; xlix, 1929, pp. 169-246; lii, 1932, pp. 105-176, 215-231; liii, 1933, pp. 170-187. ———.— Mitt. Zool. Mus. Hamburg, xliii, 1928, pp. 155-198. Bull. Mus. Hist. Nat. Paris, (2), iv, 1932, 872-881. —_—_———.—-Stett. Ht. Zeit., xciii, 1932, pp. 220-241. Ann. Mag. Nat. Hist., (10), xiii, 1934, pp. 1-38. ADDITIONS TO OUR KNOWLEDGE OF THE FLORA OF THE NARRABEEN STAGE OF THE HAWKESBURY SERIES IN NEW SOUTH WALES. By N. A. Burress, M.Sc. (Plate x; eleven Text-figures. ) [Read 28th August, 1935.] During 1933 a systematic examination was undertaken by the Students’ Geological Society of the University of Sydney of the fossil flora of portions of the Narrabeen Stage. This Stage is as yet very imperfectly known, notwithstanding its ready accessibility from Sydney. Plant remains and geological data were collected. The Research Committee of the Society consisted of its President, Mr. M. D. Garretty (Chairman), Mr. N. A. Burges, M.Sc., of the botanical staff at the University of Sydney, and Mr. S. W. Carey, B.Se. The following members participated in the excursions which were held in addition to frequent committee visits: Misses T. Christie, M. Cogle, G. Edgecombe, M. Hayward, B. E. Johnston, V. M. B. May, D. Pearce, N. Repin, N. Robards, EH. J. Thompson, N. L. Wilkie; Messrs. F. A. Hanlon, D. J. Lee, R. McGlynn, K. MacKinnon, W. McNiven, K. Mosher, W. Nichols, L. Noakes, J. Pryke, M. L. Wade, J. Yager. Thanks are due to Dr. A. B. Walkom for much help in connexion with the plant determinations, and also to Professor T. G. B. Osborn, and to Professor L. A. Cotton, for permission to carry out work in the Departments of Botany and Geology, respectively, in the University of Sydney. The geological notes on the Narrabeen Stage are due to Mr. M. D. Garretty. GEOLOGICAL SUMMARY. The Triassic Hawkesbury Series of the Sydney District has been divided into three stages, the lowest being the Narrabeen Stage. This is for the most part conformable with the underlying Kamilaroi (Permo-Carboniferous) Coal Measures; it is overlain conformably by the Hawkesbury Sandstone Stage. It was placed by the late Sir T. W. Edgeworth David (1932) as early Triassic (Lower Bunter), and the beds were tentatively regarded as Lower Triassic on botanical grounds by Walkom (1918). Fossil fish and a labyrinthodont lead to the same conclusion. Unio indicates the freshwater nature of the sediments. The palaeontological equivalent of the structural continuity of Palaeozoic and Mesozoic exists locally as a gradation in the flora for a few feet (Dun, 1910). The Stage forms part of the Sydney Geosyncline, and has at Sydney a maximum thickness of about 1,800 feet. It thins out to the south (about Kiama), west (about Lithgow), and north (Gunnedah and Murrurundi), and is truncated to the east at the coastline. Outcrops are in general restricted to the coast and margin of the geosyncline. The sediments near Sydney consist largely of shales and carbonaceous shales, with associated sandstones, tuffs, and hard L 258 FLORA OF THE NARRABEEN STAGE, N.S.W., fontainebleau sandstone in thin bands. To the north, west, and south, coarser types prevail. On the South Coast near Otford numerous pebbles apparently derived from a Narrabeen conglomerate were found; these in part closely resemble certain Upper Devonian lavas from Yalwal (50-60 miles to the south- south-west). Fragments of large trees and smaller plant remains occur here in a charred state in the tuffs, indicating proximity to points of eruption. These occurrences strengthen the view (David, 1887) that “the probable source of these [cupriferous, see below] particles is the line of volcanic country between Kiama and Mittagong”. A kind of rhythmical alternation of sandstone and shale, on various scales, is not uncommon. The coarser bands frequently have macerated plant remains showing internal signs of far-carriage, while much better preserved specimens occur in the finer laminae. Important and distinctive beds are the HMstheria Shales, Cupriferous Tuffs, and the Chocolate Shales. The first (with H. coghlani) overlie the Kamilaroi sediments for about 560 feet, below Sydney. Copper-bearing tuffs follow for about 40 feet at Sydney, and also occur higher in the Stage, associated with Chocolate Shales. The latter form a band up to 170 feet thick near Sydney, near the top of the Stage; elsewhere they thin out, and may split into several bands. The Chocolate Shale is considered to be a redistributed tuff, with admixed sediment. The top of the upper Chocolate Shale zone is sometimes taken “for convenience” (Harper, 1915) as the actual top of the Narrabeen Stage, but this is not the case, as it may continue for some distance higher up, as at Sydney. This is important, since most of the plants described were collected not far above the upper limit of the chocolate shale, the principal localities being: Turrimetta Head, Mona Vale, Avalon, and Terrigal. (For a sketch geological map of the area see Culey, 1932.) Recent work (Culey, 1932) on ripple-marks has shown “the Triassic Narrabeen Lake as a shallow, subsiding, freshwater lake, probably elongated in a N.H.-S.W. direction. Surrounding it one would see areas of low relief from which the sediments are brought down and deposited quietly in the lake, the prevailing calm being interrupted by local disturbances and ejections of tuffs followed again by quiet sedimentation”. Washaways are common in sectional form in the cliffs and could be due to sub-aerial erosion, or to local channels in sheet flood erosion. There is abundant evidence of deposition in separate hollows or lakes, and the beds are lenticular (in extent measurable usually as a fraction of a mile). For this reason the Society’s original project of zoning the Stage by its flora is seen to be impracticable. Periodical floods seem to have brought sediment and plants for some distance, forming persistent pebble bands and torrential bedding, accompanied by plant remains in a macerated condition. At quieter times plants would come merely from the borders of the individual ponds, and be gently covered bv fine sediment. That at times part of the area was dry land is indicated by the occurrence in interformational conglomerates (as at Mona Vale immediately above the Chocolate Shale) of fragments of already solidified shale from elsewhere in the Stage; one such fragment contains a well-preserved Cladophlebis. The work of this paper, geological and botanical, has thus given to the Narrabeen Landscape a greater heterogeneity than was formerly supposed. While occasional events of an all-embracing nature took place, the more peaceful times saw semi-isolated and moisture-zoned plant communities around the separate small lakes, resembling numerous oases in a desert of what was probably dry land (plain of accumulation). BY N. A. BURGES. 259 Certain structures were described by Walkom (1925, plate xxxi, figs. 7-9) as “Plantae incertae sedis”. Of these, figs. 7 and 8 have been referred to Araucarites sydneyensis (q.v.). Occurrences of the structure represented by fig. 9 were found to be quite numerous on certain horizons at Turrimetta Head. They were found not to be referable to a plant or plants, but to be concretions. They are generally dome-shaped, with surface, and the cavity of the overlying shale, striated radially and having a slickensided appearance. Slides show that the concretions have formed in bands of shale with an unusually fine texture; no nucleus was observed as such. Application of the criteria of Richardson (1921) did not lead to decisive classification, but there seems no doubt in this case that the concretions are subsequent rather than contemporaneous. Nodules of an apparently similar type, but without the slickensides, have been collected in the Triassic Wianamatta Stage at Strathfield. PLANT DESCRIPTIONS. The following section of the paper contains descriptions of plants found during the course of the work. Some record additional features for species already described, while others are new species. Well-known plants previously recorded from the Narrabeen stage are not discussed, as these have been treated fully by Walkom (1925). The distribution of the plants is interesting—seldom were many types found on one horizon. Usually the beds contained Thinnfeldia and Phyllotheca almost to the exclusion of other plants, but here and there beds occur which contain little or no Thinnfeldia but are rich in other plants. At Mona Vale an horizon was found in which all the plant remains were referable to Araucarites; at Avalon extensive beds were found with Williamsonia stems. Of the following records, the occurrence of fossil wood of the cupressinoid type from the base of the Triassic is of most interest. LYCOPODIALES. LYCOSTROBUS LONGICAULIS, n. sp. Text-fig. 1. Included in the material collected from Avalon are several specimens of cones. These structures are borne on stems which in some cases are a metre in length. The stems are 2 cm. wide and very long, they show no trace of structure other than a few longitudinal striations, some of which are probably due to folding during preservation. The cone itself is about 8 cm. long by 4 cm. wide and tapers away towards the apex. It resembles fairly closely in general appearance a male cone of Macrozamia spiralis even to the suggestion of pointed sporophyls with upturned ends, becoming longer towards the apex of the cone. The sporophylls are spirally arranged. Little can be made of their structure. The specimens are partly casts and partly composed of organic matter. Treatment with hydrofluoric acid followed by nitric acid and potassium chlorate shows the presence of the remains of both thick and thin walled cells and structures which are probably spores; these are much distorted but would be about 204 in diameter. FILICALES. TODITES NARRABEENENSIS, n. sp. Text-fig. 2. Fronds pinnate, probably 30-50 cm. long or more, pinnae linear, pinnatisect, at least 20 cm. long; 8-5 mm. broad in the fertile material, 5-8 mm. in the sterile; the segments in the fertile material orbicular or reduced to crenulations 260 FLORA OF THE NARRABEEN STAGE, N.S.W., of the pinna margin; segments of sterile pinnae acute, slightly falecate. Sporangia scattered over the undersurface as in 7. Williamsoni (Brong.). T. narrabeenensis differs from JT. Williamsoni in its much smaller size and its sharp distinction of the sterile from the fertile material. Material of this species is not plentiful and appears to be concentrated in two or three layers in the grey shales about forty feet above the chocolate layers. Specimens were collected from Turrimetta Head, Mona Vale and Avalon. t S a a 0 = ) dO yO 9% —~.. ») & ) Wi a = 0 “ Text-fig. 1.—Lycostrobus longicaulis. Cone and portion of stalk, x 4. Specimen No. 2003.* Text-fig. 2.—Todites narrabeenensis. a, Fertile frond, x 4; b, pinna showing sporangia, x 3; ec, sterile pinnae, x 4. Specimen No. 2004. Text-figs. 3, 4.—Caulopteris sp.? 3, Stem with expanded apex, x 4. 4, Leaf scar enlarged, x 22. Text-fig. 5.—Expanded apex of a stem probably referable to Caulopteris, x 3. Specimen No. 2006. Text-figs. 6, 6a.—Odontopteris dubia. 6, Frond, x 3. 6a, Portion enlarged, x 2. Specimen No. 2007. FERN STEMS. CAULOPTERIS sp.? Text-figs. 3, 4. In one horizon at Avalon, large numbers of casts and impressions of stems were found. Some of these are quite large—up to 5 cm. in width and 20 cm. long. The main part of the stem is covered with a series of scars, irregularly placed but tending to form rows with ridges of raised tissue between them. These ridges are usually not regular and are often oblique to the main axis of the stem. The apex is slightly expanded, bearing scars more closely set and less typical in shape. Intermediate types of scars are present at the base of the expanded * The numbers are those of specimens in the collection of the Geology Department, University of Sydney. BY N. A. BURGES. 261 portion. The stem is probably partly or wholly decorticated in the lower region. The scars are elliptical, 2-3 mm. by 15 mm. (Text-fig. 4), and show markings which are interpreted as representing an upper sclerenchyma band of tissue and a U-shaped vascular strand. The specimen illustrated in Text-figure 5 probably also belongs here. Both are referred with doubt to the genus Cauwlopteris. PTERIDOSPERMS? ? TAENIOPTERIS UNDULATA, nN. sp. Plate x, fig. 1. Leaf linear, at least 0-5 metre long, 1:4 cm. wide, midrib very prominent, about 2 mm., lamina very regularly undulate in most specimens, venation taeniopteroid, at right angles to the midrib, veins very delicate, numerous, 30 per cm. This species resembles in some ways T. spathulata, but no basal or apical parts have been seen even in pieces 40 cm. long. ODONTOPTERIS DUBIA, N. sp. Text-fig. 6, 6a. Leaves simple, linear, lanceolate, 10-12 cm. long, 2 cm. wide, pinnatisected segments rounded, venation odontopteroid. Tapering at the base. GINKGOALES.. BAIERA SiImMoNpDSI (Shirley). Text-figs. 7, 8. Several specimens are probably referable to this Queensland species. There is in one specimen (Text-fig. 8) a suggestion of a midvein in each linear segment. This may be due to folding during preservation, or may indicate that the specimen belongs elsewhere. It is certainly larger than the average Baiera Simmondsi. Text-figs. 7, 8.—Baiera Simmondsi (Shirley). 7, Portion of leaf, x 4. 8, Portion of leaf, x 4. Specimens No. 2008, 2009. Text-fig. 9.—Rhipidopsis narrabeenensis Walkom. Leaf showing petiole, x 3. Specimen No. 2012. Text-figs. 10, 11.—Araucarites sydneyensis Walkom. 10, Cross section of cone, x 3. 11, Cone scale, x 2. Specimens No. 2010, 2011. 262 FLORA OF THE NARRABEEN STAGE, N.S.W., RHIPIDOPSIS NARRABEENENSIS Walkom. ‘Text-fig. 9. Further material of the above species confirms Walkom’s (1925) opinion that it belongs to Rhipidopsis rather than Psygmophyllum. The specimen figured (Text-fig. 9) possesses a petiole about 5 cm. long—it agrees with that figured by Walkom (1925, fig. 4, Pl. xxx) in having the two segments forming the leaf blade. CONIFERALES. ARAUCARITES SYDNEYENSIS Walkom. ‘Text-figs. 10, 11. The following additions can be made to the description of the above species. The remains described by Walkom of this species are evidently those of megaspore cones. The specimens illustrated by Walkom (1925, Pl. xxxi, figs. 7, 8) and placed by him in Plantae Incertae Sedis are probably cross sections of a cone referable to this species. A large series of specimens obtained from Mona Vale shows that the cones were preserved in such a way that all aspects of the cone may be examined. The cones appeared to fracture easily and many were preserved showing cross-sections in which the scales may be seen in situ (cf. Text-fig. 11). Large numbers of detached cone scales were also found. The cone scales are approximately triangular, about 1 cm. long and about 5 mm. broad, tipped by a short point, with a marked ridge along one side. PETRIFIED MATERIAL. Previous descriptions of plants from the Narrabeen Stage have been made from. casts, impressions or carbonized remains. During the present investigation, material was obtained from Terrigal which, although unpromising in surface view, was well preserved and yielded good sections. Two trips were made to the locality, one by Mr. Noakes, in 1933, who was making a reconnaissance of the area when he found the first piece of material, and a second in 1934 when other members of the society accompanied Mr. Noakes and collected specimens of at least two species. The remains consisting of petrified stems were embedded in the tuffaceous sandstones which form the cliffs of the Skillion and the headlands immediately to the south. These cliffs in most places are difficult to climb, so the main collecting was carried out among the fallen rock. In one instance material was obtained on the cliff face. Nowhere did the plant remains seem to be at all plentiful, and three to four hours’ careful search revealed only six pieces of stem. When dug from the matrix the specimens are usually surrounded by a layer of coaly material which is more or less prismatized. Frequently they are rounded at the ends, suggesting that disintegration has occurred before preservation. Considerable crushing has occurred, but this has been irregular and areas of undisturbed material are available for examination. The preservation is due to infiltration of calcareous substances, and much of the original carbonaceous material is still present. The cellular structure is well retained and the tracheidal pitting is easily visible. In some specimens the protoxylem elements and the pith cells are recognizable. Three forms of the material are present, two of these can be assigned to Cupressinoxylon; the third is difficult to place, but is here considered as Cedroxylon. BY N. A. BURGES. 263 CUPRESSINOXYLON NOVAE-VALESIAE, n. sp. Plate x, figs. 2-5. The specimens are portions of stems or branches, and vary in length, one measuring 70 cm. Owing to compression and also disintegration, they have assumed an oval shape, 5-5 cm. by 2-8 em., with an ex-centric pith. The pith is well preserved in some pieces but in others has been lost; it measures 3-8 mm. in diameter. The cells increase in size towards the centre, 40u-60u in diameter, the walls with circular pits, air spaces between the cells triangular, 84. Protoxylem endarch, tending to be split up into wedge-shaped pieces. Tracheids nearest the pith small, 10%, annular or spiral, metaxylem larger, 30u, scalariform. Secondary wood showing somewhat irregular annual rings, 0-5-3.mm. wide; spring tracheids large, 30-40u, slightly irregular, summer tracheids smaller, 20-30u, and thicker walled, pits in a single row, bordered with circular orifice, rims of sanio apparently poorly developed, usually not distinguishable; checking is evident. Rays simple, 2-8 cells deep, usually 2-4, the cells 20 by 30u in tangential section and 50-60u long, tangential pits not clearly shown, 2-4 in the field. Proportion of medullary ray to the wood in tangential section about 1:15. Resin tissue consisting of isolated rows of parenchyma cells fairly evenly distributed, abundant. In addition to the above, there is material of Cupressinoxylon (Pl. x, fig. 6) very Similar to C. novae-valesiae but differing somewhat. The general features vary little in essential points, the differences being more apparent than real. In cross-section the protoxylem is more irregular than in C. novae-valesiae as is the secondary xylem. The only well marked feature is the size of the medullary rays, which are 6-9 cells deep and usually narrower than in C. novae-valesiae. ? CEDROXYLON TRIASSICUM, n. Sp. PI. x, figs. 7-9. Specimens are parts of stem showing fairly well preserved pith and xylem, Oval in cross-section, partly due to crushing and partly to weathering or decay prior to embedding. The pith is about 8 mm. in diameter, the cells about 40u in diameter, the walls not very definitely pitted. Protoxylem fairly abundant, in wedge-shaped pieces jutting into the pith. The smaller elements spiral, 154 in diameter, the larger scalariform, 254 in diameter. Secondary wood with well-marked annual rings 2-4 mm. apart. Tracheids 30u in diameter, the summer wood composed of slightly smaller tracheids with thicker walls, bordered pits in a single row, with oblique orifices, rims of sanio very distinct, checking well marked. Rays simple, 3-10 cells deep, usually 4-5 deep, cells 18u by 25u in tangential section, 50-60u long in radial section, walls with 2-6 pits in the field. Wood parenchyma scarce, one or two cells among the summer wood, resinous. References. CuLrey, A., 1932.—Ripple Marks in the Narrabeen Series. Journ. Roy. Soc. N.S.W., xvi, p. 248. Davip, T. W. E., 1887.—Cupriferous Tuffs of the Passage Beds between the Triassic Hawkesbury Series, and the Permo-Carboniferous coal measures of N.S.W. Rept. Aust. Assoc. Adv. Sci., i, pp. 275-290. ,1932.—Explanatory Notes to accompany a New Geological Map of the Commonwealth of Australia. Aust. Medical Publishing Co., Glebe, N.S.W. Dun, W. 8., 1910.—Notes on some fossil plants from the roof of the coal seam in the Sydney Harbour Colliery. Journ. Roy. Soc. N.S.W., xliv, p. 615. Harper, L. F., 1915.—Geology of the Southern Coalfield. Mem. Geol. Surv. N.S.W., Geol. No. 7. RICHARDSON, W. A., 1921.—Concretions. Geol. Mag., lviii, pp. 114-24. 264 FLORA OF THE NARRABEEN STAGE, N.S.W. WALKOM, A. B., 1918.—Geology of the Lower Mesozoic Rocks of Queensland. Proc. LINN. Soc. N.S.W., xliii, p. 95, ete. , 1925.—Fossil Plants from the Narrabeen Stage of the Hawkesbury Series of New South Wales. Proc. LINN. Soc. N.S.W., 1, 1925. DESCRIPTION OF PLATE X. Fig. 1.—? Taeniopteris undulata, n. sp. (Specimen No. 2005). Figs. 2-5.—Cupressinoxylon novae-valesiae, n. sp. 2, Transverse section of pith and protoxylem. 3, Secondary wood showing annual rings. 4, Tangential section of the wood. 5, Radial section of the wood showing the bordered pits. Fig. 6.—Cupressinoxylon sp. Primary wood and early secondary wood. Figs. 7-9.—Cedroxylon triassicum, n. sp. 17, Longitudinal section of the protoxylem. 8, Longitudinal section of the secondary wood. 9, Longitudinal section through a medullary ray. Pe nex. N.S.W., 1935. Soc. LINN. Proc. ‘a -s0anegy a2 Tan a GREET Se pee ta as + ) oe ays moe tO lon s y Cupressinox 6, ylon novae-valesiae ; ylon triassicuwm. WPressinox 5 G 2-5 undulata ; Taeniopteris iL. edrox 9, C ie UPPER PERMIAN INSECTS OF NEW SOUTH WALES. III. THE ORDER COPEOGNATHA. By R. J. Tirtyarp, M.A., Sc.D. (Cantab.), D.Sc. (Sydney), F.R.S., HERE ON Zee HRS) SEEGss: (With thirteen Text-figures. ) [Read 28th August, 1935.] This paper is a continuation of the series which I began in 1926 (Tillyard, 1926a) with a paper dealing with the Upper Permian fossil insects of the Order Hemiptera, and followed with a second part (Tillyard, 19260) on the Orders Mecoptera, Paramecoptera and Neuroptera. During the intervening nine years, a very large number of new fossil insects have been discovered in Upper Permian rocks from Warner’s Bay, thanks to the persistent labours of Messrs. T. H. Pincombe, M. S. Stanley, Rev. A. J. Barrett and myself; to whom must be added more recently Master Malcolm Stanley, who in the course of a year or so has discovered quite a considerable number of fine wings. The insect fauna of the Upper Permian in New South Wales, as now presented to us from a study of about five hundred specimens, is a very remarkable one for a Palaeozoic Fauna. Most of the older types of insects are either absent or very rare. Orthopteroid insects of all kinds appear to be entirely absent, including Cockroaches. No Palaeodictyoptera, Megasecoptera or Mayflies have been found, and only a single larva of definite Perlarian affinity. Dragonflies are so far represented only by two or three fragments of a wing which appears to belong to the family Ditaxineuridae, known only from the Lower Permian of Kansas. The Suborder Homoptera of the Order Hemiptera was the dominant group. As these insects must have fed mainly on Glossopteris, their remarkable abundance must be taken into account as a possible factor in the decline and eventual disappearance of the Glossopteris flora at the close of the Upper Permian. Closely related to the Homoptera, and by no means uncommon, were the members of the Order Copeognatha or Psocoptera, dealt with in this Part. Apart from the above Hemimetabolous Insects, the fauna is mainly composed of Holometabola and their ancestors. Quite a number of new and interesting types of Coleoptera have been discovered, together with further representatives of the Order Protocoleoptera. Next to the Homoptera, the most abundant Order of Insects was the Mecoptera or Scorpion-flies, of which many new types have been found. The derived Order, or Sub-order, Paratrichoptera is represented by quite a number of primitive genera, and these in turn had already produced, alongside of them, true representatives of the Order Diptera. The only other Order known for certain from these beds is the Neuroptera Planipennia, of which a number of very fine new types have been discovered. There is one wing, which is unfortunately very fragmentary, which may belong to some primitive type of Hymenopteron, but this is not at all certain. M 266 UPPER PERMIAN INSECTS OF NEW SOUTH WALES. iii, The present Part deals with the Order Copeognatha only, and presents for the first time a Copeognathous fauna of great richness and variety which is evidently a marked advance on that of the Lower Permian of Kansas. Order CoPEOGNATHA. The earliest known representatives of this Order come from the Lower Permian of Kansas, and were described by me in 1926. Two families are there represented, viz., the Dichentomidae and the Permopsocidae. So far, no Copeognatha have been described from the Upper Permian of New South Wales. But, in the more recent collections from Warner’s Bay, this Order proves to have been well represented by more than twenty specimens. Unfortunately the conditions of fossilization are generally such that the wings become badly crumpled or torn. This is particularly the case with a fine species belonging to the family Dichentomidae. I have before me several specimens of which I am quite unable to give descriptions. Not only are the wings torn or crumpled, but it is evident from the habit of this insect of resting (and, evidently, also dying) with its wings held close together in a steep roof-wise position, either all four wings, or sometimes only two, became stuck together in the glue-like mud in which they were preserved, leaving impressions of either four or two very faint systems of venation, crossing one another at slight angles, and making it impossible to restore the venation as it originally was. So far, the family Permopsocidae has not been discovered in the Upper Permian beds of Australia; unless, perhaps, the new genus Megapsocidium should happen to belong to this family, when the structure of its cubitus is revealed. The Dichentomidae are well represented, and there are also two remarkable new families possessing quite unexpected types of venation. One of these, the Zygopsocidae, would appear to be a specialized group which has left no descendants. But the other, the Zoropsocidae, proves to be of special interest for two reasons: firstly, it appears to represent the ancestral stock of the recent Suborder Zoraptera, now confined to the nests of Termites; and, secondly, it supplies a connecting link between the more typical Copeognatha and the family Lophioneuridae (inclusive of Dr. F. M. Carpenter’s family Cyphoneuridae), which can now be shown to be true Copeognatha and not Homoptera as we both originally supposed. Included in the present collection are a number of new genera allied to both Lophioneura and Cyphoneura. Some of the specimens are practically complete, and prove to be insects having the general character of Copeognatha, with depressed head, hypognathous mouth-parts, no sign of a sucking-beak, wings held roof-wise over the back, and other characters of the Order. It is therefore now necessary for me to remove the family Lophioneuridae from the Homoptera and to include it in the Copeognatha. It will further be abundantly evident that Carpenter’s family Cyphoneuridae (Lower Permian of Kansas) cannot stand, but must be merged with the Lophioneuridae. The Copeognatha of the Permian now prove to have been quite abundant in families, genera and species. They are evidently very closely allied to the Homoptera, and there can be no doubt that the two groups arose from a common stem. Apart from the very obvious differences in the shape of the head and the structure of the mouth-parts, it is possible to distinguish the wings of the Permian Copeognatha at once by the fact that Rs is always branched, whereas in the ‘Permian Homoptera Rs is always simple (except for short terminal branchlets in the abundantly-veined Prosbolidae). BY R. J. TILLYARD. 267 In the figures of this paper, except only figs. 7A and 9, which are complete insects, the wings are drawn always with apices to the. right, to facilitate comparison. Key to the Families of Permian Copeognatha. (Forewings only.) 1. Rs with four branches arranged in an anteriorly pectinate series ...............- CP Oana No eD ie 0 -CROME Oat) ONGHD CO) 6 GLOLCRO=0 HEEeC CREE BURA CREE EREREREE DS". 2 Family Zygopsocidae, n. fam. Rs with less (than four branches; not arranged as above .........1.0..c5++++sss- 2 2. M three-branched; Cu, forked or simple ...... “gle Mera doses Family Dinopsocidae Mart. (Upper Permian of Russia only.) MEWwithveither four OL CWO) DEAN CHES aie capers cose sce hae ee OTR eae cae, ea outs eaters 3 Bo | WME tower lorezwavelavsxol S (wl, sseweliscl GoogacadoooocdobcubooucneonnodoNnuDSuOaueDOUOONS 4 M two-branched; Cu, usually simple, rarely with a weak posterior branch ...... 5 4. Fork of Cu, deep and strongly arched, connected with M,_, above by a cross-vein SF Cherie eae CRON ONE OREO TEC, Clone ROL GUC CRs CREME ca anc Ren CESS Family Permopsocidae Till. (Lower Permian of Kansas.) Fork of Cu, less arched, often long and flat, not connected with M,, ............-. O Gr Geo ie Bon, HGF AADRG or Gep Lone G aa CuBI CHEE ene (CRO ae Tol Cee ee Re ere Family Dichentomidae Carp. (Lower Permian of Kansas and Upper Permian of Australia.) 5. Main veins more or less strongly curved, Cu, sigmoidally; Rs, M and Cu, arising SEWAMAULEN WPCA, IR, ooooccoongp0bonda Hos eooUE DOOD Family Lophioneuridae Till. (Upper Permian of Australia and Lower Permian of Kansas.) Main veins normal, Cu, not sigmoidally curved; their modes of origin also normal 510 OSOnO OEE EO. SO ORONOUD, Gib Oceana dG) PIER CeO MPIC RE einem nas pcre sear a en Family Zoropsocidae, n. fam. Family 1. DiIcHENTOMIDAE Carp. Psocidiidae, Tillyard, 1926c, p. 319.—Dichentomidae, F. M. Carpenter, 1932, p. 3. Dr. Carpenter considers the genus Psocidium Till. to be synonymous with Dichentomum Till. As the latter takes page precedence, the name of the family should be changed to Dichentomidae. Pending a restudy of the types, I think that this alteration should stand. The two genera are in any case very closely allied. There are three new generic types in the material before me, which may be distinguished as follows: 1. Medium-sized wings with the branches of Rs arising distad from the level of the end Of PEC AE WO tee R eM: et cret cuss syed ed sucks nn araiisdsucheitisnase cos len shes Haye ce maeee seeestites REL INRA AI RMR OR aR TE 2 Large eee with the branches of Rs beginning at a level well before the end of R, Le he REE Sin 8 AERP SHER EEC AGI CAT: Cor Soka ene men a men el ee Ce ee Genus Megapsocidium, n.g. 2. Se simple; R well removed from costa; M 4-branched in forewing ................ Oo 6G O20 OFO-O8O 60 O'S. 0. Cee Tee Ono OOH CCR ONT > EEE eae RC ERE IOS Roar e tons Genus Austropsocidium, n.g. Se branched; R fairly close to costa; M 5-branched in forewing .................-. Be HL OEE alo: URE UG CRE TERE CRE IS En REESE ST EEO rece eens coll are Ree re el Genus Stenopsocidium, n.g. Genus 1. AvUSTROPSOCIDIUM, n.g. Figs. 1, 2. Forewing elongate-oval in shape, the costa gently curved, the apex well rounded. Sc ending on R,. R well removed from costa. Rs either two- or three- branched, but the branches short in any case. M four-branched, normal, connected with Rs by a radio-median cross-vein. Cu, arising just below M and having a long, rather flat fork not connected with M in any way. Cu. a straight, weak furrow-vein bounding a small but well developed anal area or clavus with two anal veins; 1A nearly straight, close to Cu,; 2A slightly sigmoid. Genotype: Austropsocidium pincombei, n. sp. Horizon.—Warner’s Bay. Upper Permian of New South Wales. Key to the Species of the Genus Austropsocidium. Rs two-branched, R, without strongly marked pterostigma ...... A. pincombei, n. sp. Rs three-branched, R, with large strongly marked pterostigma .. A. stigmaticum, n. sp. 268 UPPER PERMIAN INSECTS OF NEW SOUTH WALES. ili, 1. AUSTROPSOCIDIUM PINCOMBEI, n. Sp. Fig. 1A, 1B. This specimen consists of fore and hind wings lying one upon the other, the apices being to the left. The venation of the forewing is not difficult to make out, but that of the hindwing, being fainter, is more difficult to trace. In the figures, the wings have been separated. Forewing (fig. 1, A).—Length 7:9 mm.; breadth 2-8 mm. Membrane very delicate; venation faint except for the anal veins. Se very faint basally but clearer as it approaches its junction with R,, which it does with a slight down- ward curve. R, forked distally but with little or no formation of a pterostigma. Rs arising at about one-third and forking into two just beyond the level of the end of R, M arising from R at about one-fifth and forking a little beyond middle of wing; the upper fork, M,,., is connected with Rs by a cross-vein after arching strongly upwards; its fork is at the same level as that of Rs. Fork of M,,, slightly basad from that of M,,.. Cu, running basally just below R+M and leaving it just before M, descending obliquely to fork somewhat before half-way along the wing; the long, flattish fork reaches to just beyond the level of the fork of M,,, 1A and 2A strong veins on a well formed clavus with slightly convex posterior margin. Hindwing (fig. 1, B).—Length 6-4 mm.; breadth 2-5 mm. Differs from fore- wing in the much narrower costal area and much wider space between R, and Rs. R, unbranched. Stem of M,,, much shorter than in forewing. Arch of Cu, flatter than in forewing, with Cum» placed more obliquely. Clavus smaller, apparently with only a single anal vein, 1A, present. Type.—Holotype, Specimen P. 218, found by Mr. T. H. Pincombe at Warner’s Bay, 23rd October, 1926. This was the first Psocid wing discovered in the Upper Permian beds of Australia. 2. AUSTROPSOCIDIUM STIGMATICUM, hn. Sp. Fig. 2. This species is only represented by the distal two-thirds of a rather long, slender wing, obviously a forewing, the apex to the left. Length of fragment 6-6 mm., representing a total length of about 9-5 mm. Breadth 2:6 mm. Sc not visible. R, widely forked, forming a rather long, triangular pterostigma, distinctly pigmented. Rs almost straight, forking into three short terminal branches well beyond level of pterostigma. M running parallel to Rs and forking rather narrowly below level of pterostigma; upper branch, M,,., continues almost parallel to Rs and connected with it by a perpendicular cross- vein, rm, before the fork; lower branch, M,,,, diverging slightly and forking just before level of end of pterostigma. Cu, forking only slightly before level of fork of M, and not so long or flat as in A. pincombei. Clavus and anal veins missing. Type.—Holotype, Specimen found by Rev. A. J. Barrett at Warner’s Bay in 1930. Genus 2. MEGAPSOCIDIUM, n.g. Fig. 3. Differs from Austropsocidium in having R, and Rs connected by a cross-vein below the fork of R, and in having Rs forking strongly at this cross-vein and again forked distally on the upper branch. Only the distal third of the wing is preserved, but it appears as if the four branches of M were not normally arranged, there being three branches on M,,, and M,,, being unbranched. Genotype, Megapsocidium australe, n. sp. Horizon.—Warner’s Bay. Upper Permian of New South Wales. BY R. J. TILLYARD. 269 3. MEGAPSOCIDIUM AUSTRALE, nh. Sp. Fig. 3. This specimen consists of the distal half only of a forewing (apex to the right), measuring 6:0 mm. long by 3-5 mm. wide, and probably representing a wing of total length 10 mm. or more. The apical portion is complete, but the basal break is very irregular and there is also an irregular patch of rather large size broken away from the centre. Pra pt Bip Rots Th Cu, Cun Cur, Fig. 1.—Austropsocidium pincombei, n.g. et sp.