het hy uA oy Git aga A ta tte AG pie rail Tarts ‘eh A eaAg Site fs bf Ea PAT nae tt Aten eta Vee Oh tne Woo Pease aT AP hGh ert Oe eine tee Say it %% fist fr ue Pahl heat nb Nh eh 3 Haat e % , 4 funtok tyke t atone : ‘ AN : Wonee nd tiie hasnt yh bo ; ‘ ; , 4 MLR 4 DEL Oy na t ¥ , Me ol Day rua AT ae ott Say Win ea) greg Meh Ah ate Wat RM a AC Ris AT ASTRAL RTE ot atti wal Rn eA Wt iH) YEA been mh Ay it cw) Het At ibe Wat Shay Wincele CRO AD ae aR Pro any el yg he et ey ae At 3 1h ¥ i mopntt PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SOUTH WALES VOLUME 103 (Nos 453-456; for 1978) Sydney The Linnean Society of New South Wales 1979 Contents of Proceedings, Volume 103 PART 1 (No. 453) (Issued 15th August, 1979) ANDERSON, D. T. Sir William Macleay Memorial Lecture 1978. Natunaliblistonystodayun: ss rrtteaand ccna ra tel wa canta ec ata i 1 HERBST, R. Review of the Australian fossil Dipteridaceae ............. 7 MARTIN, A. A., WATSON, G. F., GARTSIDE, D. F., LITTLEJOHN, M. J., and LOFTus-HILLS, J. J. A new species of the Litorza peronz complex (Anura: Hylidae) from eastern Australia .............. 23 DYNE, G. R. A new species of Microscolex (Diplotrema) (Annelida: Oligochaeta) from New South Wales ............... 37 FAIN, A., and DOMROwW, R. The family of Hypoderidae (Acari) PIMOS CL AT ARNE sheers pave bses tes as seae ys Vayda Gln any eas cutie wok a Lanna 43 HOLMES, W. B. K., and AsH, S. R. Anearly Triassic megafossil flora from the Lorne Basin, New South Wales .................. 47 RUSSELL, R. C., DEBENHAM, M. C., and LEE, D. J. A natural habitat of the insect pathogenic fungus Culzcenomyces in the Sydney area ... 71 THE LINNEAN SOCIETY OF NEW SOUTH WALES. Record of the Annual General Meeting 1978. Reports and Balance Sheets ....... Us PART 2 (No. 454) (Issued 15th August, 1979) WEBBY, B. D. Presidential Address 1978. The Ordovician SELOMALOPOTOIGS siya sh ee teen Roe CTS aaa sa Gre meas Fs oheiias 83 STANISIC, J. Freshwater sponges from the Northern Territory (orferaSpongillidae) se Cis see aan ie cae 123 MEMORIAL SERIES No. 26. Hans Laurits Jensen, 1898-1977 .......... 131 THE LINNEAN SOCIETY OF NEW SOUTH WALES, A key to the MemorraliSentes: (928-1078) nib. 4s teak Peet gs in tes Aes aes 133 PART 3 (No. 455) (Issued 28th December, 1979) ENGEL, B. A. Fenestrate bryozoans with large apertural form in the Carboniferous of eastern Australia ................ 135 SHORT, J. R. T. The final larval instar of Phaenocarpa (Asobara) persimilis Papp (Hymenoptera, Braconidae, Alysiinae) from Australia ..... 171 PERCIVAL, I. G. Late Ordovician articulate brachiopods from Gunningbland, central western New South Wales .......... 175 PART 4 (No. 456) (Issued 28th December, 1979) DOMROW, R. Some dermanyssid mites (Acari) , mostly from Australasiamrodents yi yin chs see ciel cic sate A Cpe ey eT ee eae 189 GEMMELL, P. Feeding habits and structure of the gut of the Australian freshwater prawn Paratya australiensis Kemp (Crustacea .CarideacAtyidae) \ 2s Serene ain eee 209 SCHICHA, E. Three new species of Amblysezus Berlese (Acarina: Phytoseiidae) from Australia .................-... 217 WATSON, J. E. Biota of a temperate shallow water reef ............. Zoi WEBB, J. A., and BELL, G. D. A new species of Lzmnadza (Crustacea: Conchostraca) from the Granite Belt in southern Queensland and northern New South Wales .................. 237 BASS, A. J. Records of little-known sharks from Australian waters ..... 247 Woop, A. E. A key to the Australian genera of the Agaricales ........ 255 DANIELS, G. A new species of Dakznomyza from Queensland (Dipterai: Asilidae) iii. se eee eee Zino TINDER Se ns Seas ROO REF RI aE SI) PICEA eg UC 283 PROCEEDINGS of the . LINNEAN | SOCIETY NEW SOUTH WALES» VOLUME 103 PARTS 1&2 ly > Ee HG AN » oe a fan i a Es Z LE) eat mt mt a oe | EG In He Dy 76 Zant UU Aye i uy Utd TE iy NATURAL HISTORY IN ALL ITS BRANCHES THE LINNEAN SOCIETY OF NEW SOUTH WALES Founded 1874. Incorporated 1884. The Society exists to promote ‘the Cultivation and Study of the Science of Natural History in all its Branches’. It holds meetings and field excursions, offers annually a Linnean Macleay Fellowship for research, contributes to the stipend of the Linnean Macleay Lecturer in Microbiology at the University of Sydney, and publishes the Proceedings. Meetings include that for the Sir William Macleay Memorial Lecture, delivered biennially by a person eminent in some branch of Natural Science. The Society’s extensive library is housed at the Science Centre in Sydney. Membership enquiries should be addressed in the first instance to the Secretary. Candidates for election to the Society must be recommended by two members. The present annual subscription is $17.50. -~ The current rate of subscription to the Proceedings for non-members is set at $27.50 per volume (including postage). Back issues of all but a few volumes and parts of the Proceedzngs are available for purchase. A price list will be supplied on application to the Secretary. OFFICERS AND COUNCIL 1978-79 President: J.T. WATERHOUSE Vice-Prestdents: D. W. EDWARDS', LYNETTE A. MOFFAT, T. G. VALLANCE, B.D. WEBBY Honorary Treasurer: JOYCE W. VICKERY’, D. A. ADAMSON? Secretary: BARBARA STODDARD Council: D. A. ADAMSON, D. W. EDWARDS’, A. E. GREER, L. A. S. JOHNSON, HELENE A. MARTIN, LYNETTE A. MOFFAT, P: MYERSCOUGH, A. RITCHIE, A. N. RODD, F. W. E. ROWE, E. J. SELBY*, C. N. SMITHERS, P. J. STANBURY, N. G.“STEPHENSON, TI.) G: VAELANGE JOVEGE OW - VICKERY, J. T. WATERHOUSE, B. D. WEBBY, A. J. T. WRIGHT Honorary Editor: T. G. VALLANCE — Department of Geology & Geophysics, University of Sydney, Australia, 2006. Librarian: PAULINE G. MILLS Linnean Macleay Fellow: JENNIFER M. E. ANDERSON Linnean Macleay Lecturer in Microbiology: K.-Y. CHO Auditors: W. SINCLAIR & Co. The office of the Society is in the Science Centre, 35-43 Clarence Street, Sydney, N.S.W., Australia, 2000. Telephone (02) 290 1612. © Linnean Society of New South Wales 1 resigned 20 September 1978 ? resigned 23 August 1978 appointed 6 September 1978 * appointed 22 November 1978 3 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 103 PART 1 THE SIR WILLIAM MACLEAY MEMORIAL LECTURE 1978 Natural History Today D. T. ANDERSON, F.R.S. School of Biological Sciences, University of Sydney [Delivered 16 March 1978] The Sir William Macleay Memorial Lecture is an occasion on which to honour the memory and legacy of one of the most important pioneers of scientific endeavour in the State of New South Wales and in Australia generally. Prime architect and benefactor of the Linnean Society of New South Wales, Sir William Macleay was foremost a naturalist, at a time when that term was a respectable admission both for a gentleman and for a man of science. Arriving in New South Wales from England in 1839 at the age of 19, he had already spent some time as a medical student, but opportunities and family associations were to divert him wholly from this course (Walkom, 1942). He became quickly involved in the management of property on the Murrumbidgee near Wagga, where he lived almost permanently for 15 years. It was from this formative period of direct and intimate contact with the flora and fauna of the Australian bush, that William Macleay developed a devotion and a commitment to the study of nature that was to dominate the rest of his life. He returned to residence in Sydney in 1857, and to comfortable circumstances which allowed him in 1862, at the age of 42, to make the decision to give his whole attention to Natural History. Out of this decision arose many investigations, expeditions and benefactions in the study of the biological sciences generally in Australia, all of which retain their importance even after the passage of nearly 120 years. Among them was the formation of the Linnean Society of New South Wales, in 1874. In initiating this move, in association with a number of other zoologists, botanists and geologists of the day, Macleay called for the formation of “A Society of Natural History”. Ten years later, in a Presidential Address to the then well established Society (Fletcher, 1893), he included the following comment: “Our rules state that the Society is for the promotion of the study of Natural History in all its branches”. In the course of time the objectives of the Society were redefined as the promotion of the study of Natural Science, but the practice of the Society has been to retain the spirit of Sir William’s major intention, in fostering the development of research and scholarship in Zoology, Botany and Geology. During the twentieth century, with the increasing penetration of science into more and more esoteric concepts and technologies, these disciplines have broadened and become cross-linked, and the Society has now come to embrace the Biological Sciences and Earth Sciences as its domain — a move which Sir William would have thoroughly approved. I feel, however, that because Sir William came to be a scientist and a promoter of scientific endeavour through his observation and appreciation of nature, and not through any formal early training, he might also have been concerned to see that the focal point of his view of Natural History, namely the continued comprehension of the faunal, floral and geological diversity of the earth, was not lost to sight in the search for ever more powerful theoretical abstractions. For example, with his interest in bacteriology (commemorated, for instance, in the Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 2 NATURAL HISTORY TODAY Linnean Macleay Lectureship in Microbiology at the University of Sydney), Sir William would no doubt have been delighted with the discoveries of modern microbial genetics; but would also have seen, I think, that these do not assist our ability to conceptualize the diversity and biology of, say, the marsupials. To do this properly one needs, among other things, a vast array of comparative descriptive and experimental data — on the animals themselves, their habits, their reproduction, their interactions with other animals and with many species of plants, their distribution, their fossil history, and the relationship of this history with the geological history of the earth in the Mesozoic and Tertiary. The same argument applies to any other group of animals or to any group of plants, with varying degrees of difficulty depending on the group. This kind of science, which focuses on the systematization of diversity, was what people meant in the nineteenth century when they talked about Natural History. Its importance was obvious at that time, because of the need to try to systematize and manage the flood of information on diversity being gathered from all parts of the world. Gradually everything fell into place with the acceptance of the idea of Evolution, which provided a means of dealing with diversity in a unified way. But from this in turn has stemmed in the present century an array of astonishing achievements in experimental biology, which has brought us to the brink of control over the fundamental processes of life, and to the powerful theoretical constructs of molecular biology and population biology. What possible significance, then, can Natural History continue to have today? Can it continue to contribute significantly to understanding, or is it a Dodo, a relic from the past that has now been knocked on the head and made extinct as a profitable, professional scientific discipline. Modern dictionary definitions of Natural History are not encouraging. Natural History, says the Shorter Oxford Dictionary (1959 edition), is the systematic study of all natural objects, animal, vegetable and mineral — so far, so good — but, now restricted to the study of animal life, usually in a popular manner. Example, another incident in natural history is, “Toads eat larks”’. Websters Dictionary (1926 edition) is even more interestingly denigrating. Natural History, it says, was formerly the study, description and classification of animals, plants, minerals and other natural objects, thus including the sciences of zoology, botany, mineralogy, etc. in so far as they existed at that time (the time is not specified, but the implication is that of Aristotle and Pliny), but now commonly restricted to a study of these subjects in a more or less superficial way, at least without making use of modern anatomical and analytical methods. A DODO INDEED! In fact, a more constructive definition of scientific Natural History can be developed by contemplating the thoughts and work of some of its founders as contributing scientists. I present two examples, both Englishmen, Gilbert White in the eighteenth century and Charles Darwin in the nineteenth century. Gilbert White, although he wrote only one book in his entire life (White, 1788), is one of the most interesting characters in the history of Zoology. He was born in 1720. His father was a barrister and his mother was the daughter of the then vicar of Selborne, a small village in rural Hampshire in southern England. Gilbert himself, after pursuing a liberal education and taking an M.A. at Oxford in 1746 at the age of 26, entered the church. For some 15 years he moved among different parishes in England, to become eventually in 1761, at the age of 40, a curate in the village of his birth, Selborne. Here he remained until his death at 72. The Selborne Parish Register records that the Reverend Mr White officiated on June 10th, 1793, at the burial of one Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 D. T. ANDERSON 3 of his parishioners, a 16-year-old girl. During this sad duty, White caught a cold, to which he succumbed before the end of the month. In the 32 years of his settled curacy at Selborne, Gilbert White became, like many educated men of his time, an ardent naturalist. His uniqueness in this pursuit lies in the manner of his investigations and in the book that he wrote on his observations, “The Natural History of Selborne”. White spent more time on Natural History than he did on the business of his parish. He identified and described many species hitherto unrecognized, thus contributing directly to the major academic activity of the zoology of the day, dominated by none other than Linnaeus himself. But he also, and this is the crucial point, observed, recorded and interpreted the daily lives and activities of these species. White’s are among the first accurate and constructive observations on such matters as — the species specificity of bird songs and their use in distinguishing closely related species; the functional significance of animal colouration; the phenomenon of territoriality in birds; the phenomenon of bird migrations; the occurrence of pseudo-copulation of frogs; and the nocturnal activities of bats, large and small. Never formally trained in zoology, he set in motion an emphasis on field studies and the observation of living animals, of which James Fisher, in his introduction to the 1947 edition of The Natural History of Selborne (Fisher, 1947), had this to say: “The theory of organic evolution could never have been propounded, as it was, in the middle of the nineteenth century, without two centuries or more of serious classification, and fifty years or more of serious field observation. It was useless to know how animals were built without knowing the quality of their lives”. Gilbert White investigated the quality of their lives. He wrote, for example, of the night-jar Caprimulgus feeding on chafers at an oak tree, that “I saw it distinctly, more than once, put out its short leg while on the wing, and, by a bend of the head, deliver somewhat into its mouth. If it takes any part of its prey with its foot, as I have now the greatest reason to suppose it does these chafers, I no longer wonder at the use of its middle toe, which is curiously furnished with a serrated claw”. The approach is modern — field observation, cautious interpretation, emphasis on living function. In fact, the claw now appears to be more important in preening — but at least White based his remarks on an observation of nature and not on an imaginative interpretation of dead specimens. Not that he was loath to investigate a dead specimen in connection with his studies if necessary. Take, for example, these comments on a species of large bat. “This summer through I have seen but two of that large species —— ; I procured one of them and found it to be a male; and made no doubt, as they accompanied together, that the other was a female; but happening in an evening or two to procure the other likewise, I was somewhat disappointed when it appeared to be of the same sex — amply furnished with the parts of generation, much resembling those of a boar.” Here is evidence of detailed anatomical as well as field observation. White was, in fact, the epitome of the well rounded, eighteenth century clerical gentleman. He even adhered to the tradition of the day by writing poetry, though alas without the perceptiveness and discrimination that he applied to his observations of Nature: “Ts this the scene that late with rapture rang, Where Delphy danced, and gentle Anna sang ; With fairy step, where Harriet tripped so late, And on her stump reclined the musing Kitty sate?” The curate’s calling is more evident here, in a portion of an ode on a visit by three eligible sisters to his bachelor field haunts. It is perhaps no wonder that Gilbert remained a bachelor. Nevertheless, he understood the meaning and purpose of Natural History in a remarkable way. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 4 NATURAL HISTORY TODAY The inspiration engendered by the observational techniques of the Reverend Gilbert White soon encouraged a deeper investigation of animals in the field rather than in museums. From about 1820, one begins to see a vast increase in interest in Natural History as recorded from field observations, in a period that was to last through much of the century and was to see the birth of the Theory of Evolution as a consequence. There is no doubt that the unparalleled contributions to Biology made by Charles Darwin had their origin, among other things, in the experiences he gained during his participation in the five years voyage of the Beagle. In the preface to his remarkable account of these experiences, the “Voyage of the Beagle”, Darwin (1845) directs attention to the following matters: 1. That the volume contains a history of the voyage and a sketch of those observations in Natural History and Geology which possess some interest to the general reader. 2. That in a larger publication, the Zoology of the Voyage, he had appended to the description of each species (described systematically by various specialists of the day) an account of its habits and range. It was this awareness of habits and range, that is, of observations on the lives of species in their natural habitat, and the relating of these observations to structure and adaptation, which gave all of Darwin’s work its modern feel and which led him with great insight to the possibility of a system of generalization through which all such phenomena might be comprehended. He was supremely aware of the diversity, functional interrelatedness and temporal instability of living phenomena, and of the relationships between those endless minutiae of detail concerning animals and plants that can be expressed verbally and graphically, but cannot be reduced to mathematical or chemical formulations. And he could express this awareness with the most penetrating simplicity: “The slimy, disgusting Holothuriae, which the Chinese gourmands are so fond of, also feed largely on corals; and the bony apparatus within their bodies seems well adapted to this end. These Holothuriae, the fish, the numerous burrowing shells, and nereidous worms, which perforate every block of dead coral, must be very efficient agents in producing the fine white mud which lies at the bottom and on the shores of the lagoon” (Voyage of the Beagle, p. 463, Keeling Island). And again: “It was most striking to be surrounded by new birds, new reptiles, new shells, new insects, new plants, and yet by innumerable trifling details of structure, and even by the tones of voice and plumage of the birds, to have the temperate plains of Patagonia, or the hot dry deserts of Northern Chile, vividly brought before my eyes. Why, on these small points of land, which within a late geological period must be covered by the ocean, which are formed of basaltic lava, and therefore different in geological character from the American continent, and which are placed under a peculiar climate — why were their aboriginal inhabitants, associated, I may add, in different proportions in both kind and number from those on the continent, and therefore acting on each other in a different manner — why were they created on American types of organization?” (Voyage of the Beagle, p. 393, Galapagos Archipelago) . One can perhaps sum up and define Natural History as Darwin understood it in the following way. It is the investigation of: the diversity of animal and plant life, the relation of structure to habit and environment, the perpetuation of diversity through reproduction and the evolution of diversity through time and the distillation from these observations of generalizations which summate this diversity. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 D. T. ANDERSON 5 Using this definition, we can now address the question, whether Natural History has a role today. In my opinion it has, and for the following reasons: Our knowledge of diversity has increased enormously in the twentieth century. Vast numbers of new species have been identified and named. The amount of information on the distribution, structure, functional organization, activities, reproduction and life cycles of known species has increased to a torrent. A similar plethora of information has been gathered on fossil species, especially of invertebrates, and new techniques have permitted fossil material to be interpreted in much more detail with respect to modes of life, interrelationships and temporal distribution and spatial distribution. All of this information needs to be incorporated into new generalizations which convey our present level of understanding, and not simply tacked on to generalizations which were propounded in the nineteenth century. Let me trace through an example with which I am familiar, progress in the understandings of barnacles. Darwin’s two classic monographs (1851, 1854), systematized all there was to know about barnacles at that time and provided a comprehensive statement about the phylogeny, evolution and biology of this group. Subsequent studies have modified this statement in various directions, concerned with feeding, reproduction, embryonic development, larval development, population biology and fossil history. With the exception of the latter, however, these studies have been conducted in relation to various kinds of experimental conceptualizations in physiology, ecology and developmental biology. Only the palaeontologists have kept in mind that questions about barnacles can be most usefully framed in the context of a comprehensive overview of the group, and have tried to improve on Darwin’s overview in the light of new information. This endeavour has now culminated in an extensive revision by Newman and Ross (1976) of the phylogeny and classification of the balanomorph barnacles, based on recent and fossil skeletal structure; but much of the other information gathered on structure, function and reproduction in barnacles during the last 120 years still lies outside this framework. It is easy to see what happens as a result. The natural history of barnacles continues to be expressed in outmoded terms, and erroneous conceptions of our basic knowledge of these animals continue to be incorporated into otherwise sophisticated physiological, developmental and ecological investigations. Suppose, for example, that one wishes to investigate an ecological problem involving knowledge of the pattern and composition of food intake by a particular species of barnacle. Is the information available? Probably not. In order to obtain it, what does one have to use? The techniques of natural history. In order to apply these successfully, what does one have to know? How to describe and interpret one’s observations in the general context of information about feeding mechanisms and their evolution in the Cirripedia. Where can this information be found? In Darwin (1854) ; and then in a random scatter of observations that have never been correlated one with another or used to modify, as they must, Darwin’s initial interpretation. We could do better. As was so eloquently argued by J. W. Evans (1965), we should do better. We should encourage the improvement of comparative, whole- organism descriptive and experimental biology (Natural History) all the time, by professional biologists for professional biologists, so that we can continue to come to terms with diversity as well as with causality. Both are aspects of the same truth. In fact, I would be prepared to go one step further and plead the cause of professional scientific Natural History in its contribution to the larger human endeavour. The conflict between exploitation and conservation must be resolved if we are not all to sink into the mire of a murdered world. We, as biologists, have a prime responsibility in promoting the conservation of nature. Part of this responsibility can Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 6 NATURAL HISTORY TODAY be met by professional scientific work leading to suitable management techniques — but this work is often highly mathematical and always too high powered for the average person to grasp. Yet the community at large has to be persuaded, because it eventually has to foot the bill in limiting exploitation and accepting that conservation has advantages. Education in Natural History, which builds on a natural link between man and nature, which presents the familiar, shows it to be complex and worthy of respect, and puts this complexity in visual, verbal, graspable terms, can provide one of the few means of communication that can prevent the community from becoming disenchanted with science. References Darwin, C., 1845.— The Voyage of the Beagle. Natural History Library Edition, annotated and with an introduction by Leonard Engel, Anchor Books, Doubleday and Company, Inc., New York, 1962. ——, 1851.— A monograph on the subclass Cirripedia. 1. The Lepadidae or pedunculate cirripedes. Ray Society, London. ——, 1854.— A monograph on the subclass Cirripedia. 2. The Balanidae, Verrucidae, etc. Ray Society, London. Evans, J. W., 1965. — The future of Natural History. Aust. J. Scz., 28: 105-112. FIsHER, J., 1947.— Introduction, pp. ix-xxi. In The Natural History of Selborne, 1947 edition, Cresset Press, London. FLETCHER, J. J., 1893.— The Hon. Sir William Macleay, Kt., F.L.S.. M.L.C. The Macleay Memorial Volume, ed. J. J. Fletcher, Linnean Society of New South Wales, Sydney. NeEwMaN, W. A., and Ross, A., 1976. — Revision of the balanomorph barnacles; including a catalogue of the species. Mem. San. Diego Soc. Nat. Hist., 9: 1-108. WaLkono, A. B., 1942.— The background to Sir William Macleay’s endowment of Natural History. Proc. Linn. Soc. N.S.W., 47: 6-15. WuiteE, G., 1788.— The Natural History of Selborne. Edited and with an introduction by James Fisher. Cresset Press, London, 1947. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 Review of the Australian Dipteridaceae RAFAEL HERBST (Communicated by J. F. RIGBY) HERBST, R. Review of the Australian Dipteridaceae. Proc. Linn. Soc. N.S.W. 103 (1), (1978) 1979: 7-21. A critical examination is made of the available fossil fern specimens included in the family Dipteridaceae from Australian collections. All valid fossil species from Australia are reviewed, but extant species are not considered. Poorly-described species are redescribed, and illustrated, with lectotypes selected where necessary. Thaumatopteris shirley sp. nov. is erected, being the first record of the genus from Australia; doubtfully, Clathropterzs is cited for the first time. The species Dictyophyllum rugosum, D. obtusilobum? and Hausmannia buchi are shown to be absent from Australia. Rafael Herbst, Faculty of Sciences, 9 de Julio 1449, Corrientes, Argentina; manuscript recetved 11 October 1977. INTRODUCTION Other than in South America, the Gondwana record of fossil members of the fern family Dipteridaceae seems to be a meagre one. Very little is known of the family from South Africa and India, and only a few ill-defined species had been described many years ago from Australasia. Considering the relative wealth of species and specimens from South America this always seemed rather strange to me. Thus, while studying Triassic plants, and having found some good specimens of these ferns, I decided to review the whole group from Australia and New Zealand without restricting it to Triassic representatives. This contribution is to clarify some names, to revalidate and redescribe some poorly known but sound species and to describe the recently found Thaumatopterts shirley? sp. nov. A list of all references to fossil Dipteridaceae was compiled, and after critical examination of the specimens themselves, many were excluded and some others were re-identified. The resulting list of taxa, here considered valid, comprises seven species included in three genera, and the doubtful presence of a fourth genus. Fortunately most of the original specimens could be located in museum collections, mainly in Queensland, therefore lectotypes and paratypes could be selected and designated. I did not see New Zealand material. Repositories of specimens are abbreviated as follows: GSQ = Geological Survey of Queensland, Brisbane MUDG = Department of Geology and Palaeontology, University of Melbourne, Melbourne QM = Queensland Museum, Brisbane SUP = Department of Geology and Geophysics, University of Sydney, Sydney UQ = Department of Geology and Mineralogy, University of Queensland, Brisbane SYSTEMATIC REVIEW Previous Records of Australasian Fossil Dipteridaceae The following list is compiled from all literature available to me in which identification of actual specimens is made: Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 8 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE Original citation Source Present status 1. Dictyophyllum bremerense Shirley, 1898 D. bremerense 2. Dictyophyllum rugosum Walkom, 1917 D. bremerense 3. Dictyophyllum davidi Walkom, 1917 D. davidii 4. Dictyophyllum acutilobum Arber, 1917 D. acutilobum 5. Dictyophyllum obiusilobum? Arber, 1917 D. sp. cf. D. acutilobum 6. Hausmannia buch Walkom, 1917 not Dipteridaceae 7. 2Dictyophyllum sp. Walkom, 1919 not Dipteridaceae 8. Dictyophyllum rugosum Walkom, 1924 D. davidit 9. Hausmannia wilkinsi Walkom, 1928 H. wilkinsi 10. Dictyophyllum bremerense Jones and de Jersey, 1947 D. bremerense 11. Dictyophyllum sp. 1 Jones and de Jersey, 1947 not Dipteridaceae 12. Dictyophyllum ? sp. 2 Jones and de Jersey, 1947 not Dipteridaceae 13. Dictyophyllum ? davidi Jones, 1948 not Dipteridaceae 14. Dictyophyllum ? rugosum Hill, 1951 D. davidii 15. Dictyophyllum bremerense Derrington, 1954 D. dawdiu 16. Hausmannia ? sp. White, 1961 H. (P.) sp. cf. H. (P.) deferrarzsi 17. Hausmanma (Protorhipis) sp. Hill et al. H. (P.) sp. cf. H. (P.) deferrarusiz 18. Hausmannza sp. White, 1966 Jet, (22) Go Gin lal, (B2,) deferrarzzsiw 19. Hausmannia wilkinst White, 1967 H. sp. cf. H. wilkinsz 20. Hausmannia sp. Douglas, 1969 H. bulbaformis 21. Dictyophyllum sp. cf. D. ellenbergz White, 1969 D. bremerense 22. Hausmannia wilkinsi White, 1972 H. sp. cf. H. wilkinsa 23. Hausmannia sp. cf. H. Gould, 1974 H. (P.) sp. cf. H. (P.) deferrariszz deferrarisi 24. Hausmannza sp. Gould, 1975 H. (P.) sp. cf. H. (P.) deferrarisi 25. Hausmannia bulbaformis Douglas, 1973 H. bulbaformis From the foregoing list, apart from the misidentified species which have been placed in synonymy, the following names are rejected or excluded from the Australasian Dipteridaceae: 5. D. obtustilobum? (Arber, 1917) : this doubtful fragment most probably is a small bit of D. acutzlobum as Arber himself expressed originally. 6. H. buchi (Walkom, 1917): this is a fragment of a fossil leaf with no visible margin at all. The veins dichotomize, but do not show traces of the anastomosing network of the Dipteridaceae. It is excluded from the record. 7. ? Dictyophyllum sp. (Walkom, 1919) : this is a fragmentary specimen which does not show any veins or other detail except its outline. It is improbable that it belongs to the Dipteridaceae. 11. and 12. Dictyophyllum sp. 1 and Dictyophyllum? sp. 2 (Jones and de Jersey, 1947) : two small fragments which do not show any trace of venation and are incomplete; they are excluded from the record. 13. Dictyophyllum? davidi (Jones, 1948) : another fragmentary specimen which only very vaguely reminds one of Dictyophyllum; a mid-vein and some lateral veins can be distinguished, but the fragment more probably is a portion of Dicrozdium sp. Specimens attributed to other species are quoted either in the synonymy or in the respective discussion below, according to the degree of confidence with which they are regarded as belonging to a particular species. The following taxa stand as valid: Dictyophyllum bremerense Shirley Dictyophyllum davidii Walkom Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST Fig. 1. 1, 2— Dictyophyllum bremerense Shirley, 1898 (1) UQ F64068 x24; (2) UQ F64068 x8. 3 — Hausmannia wilkinsii Walkom, 1928 GSQ F1943b X 134. 4 — Clathropteris sp. UQ F23071 X14.5 — Dictyophyllum bremerense Shirley, 1898 SUP 20007b X1.6 — Hausmannia (Protorhipzs) sp. cf. H. (P.) deferrarzsi Feruglio, 1937 Bureau of Mineral Resources, Canberra, specimen F22699, from Helen Springs, Northern Territory (photo kindly supplied by Mrs M. E. White) <%. 7 — Diuctophyllum davidii Walkom, 1917 GSQ F165 Holotype x 3%. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 10 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE Fig. 2. 8 — Dictyophyllum bremeren shirley’ n. sp. (9) UQ F64280 x 31%; (10) No. 2995 (CTES-P se Shirley, 1898 GSQ F12041 x14. 9, 10 — Thaumatoptervs B, Argentina) <2. ll — Hausmannia wilkinsti Walkom, 1928 GSQ F8857 x2%. 12 — Dictyophyllum davidii Walkom UQ F48643 X 2. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST 1] Dictyophyllum acutilobum (Braun) Schenk Thaumatopteris shirleyi n. sp. Hausmannia (Hausmannia ?) wilkinsa Walkom Hausmannia (Protorhipis) sp. cf. H. (P.) deferrariisiz Feruglio Hausmannia (Hausmannia) bulbaformis Douglas cf. Clathropterts sp. SYSTEMATIC PALAEOBOTANY Dictyophyllum bremerense Shirley, 1898 Hell 2,o;-big. 2-8; Fig. 3. 1-8. 1898, Dictyophyllum bremerense Shirley, Geol. Surv. Queensl., Publ. 128: 25, pl. 13, figs 2a/b. 1917, Dictyophyllum rugosum Walkom, Geol. Surv. Queensl., Publ. 257: 9 pl. 4, fig: 3; pl. 6, fig. 4B; pl. 9, fig. 3. 1947, Dictyophyllum bremerense Shirley, zn Jones & de Jersey, Univ. Queensland., Pap. Dept. Geol. 3(3) : 13; pl. 1, fig. 6; pl. 4, fig. 1b. Description: Frond palmate (?) with at least seven large pinnae united at the base, and free apically. Free parts of pinnae up to 100 mm long, probably longer, 50 mm wide, rather pinnatifid. Midrib of pinnae strong and straight; second order lateral veins (the midvein of each “pinnule”) arising at 60°, slightly falcate (or apically, concavely arched), reaching the apex of each “pinnule”’. “Pinnules” generally falcate, their free portions up to 20 mm long, by 8-9 mm wide basally; apex is generally acute, both margins strongly curved. Distance along midrib between where lateral, second order veins arise is variable, from 6-7 mm to 15- 17 mm. Third order veins (second order lateral veins) arise at 60°-70°, then by successive dichotomous divisions form a network of somewhat elongated, rectangular to polygonal meshes. The highest order meshes are 4-5 mm diameter, and, when elongated, their longest axis lies parallel to the second order veins. From the second order veins, smaller ones of successively higher orders are given off, which themselves form smaller meshes of successively higher orders. The meshes are polygonal, 4-7 sided, with the smallest, highest order, measuring about 0.5 mm diameter. Most higher order meshes are elongated, and tend to be arranged with their longest axis parallel to the second order veins. Venation is similar throughout the lamina. Fertile specimens are rare, but some show sporangia arranged in “sori” (?) either along lateral veins of second order as elongated bodies 7-8 mm long and 1 mm wide (Fig. 3. 4) or/and distributed randomly on the lamina as oval or rounded bodies 2- 3 mm diameter (Fig 3. 4, 7). The shape of these “sori” (Fig. 3. 5) is not clear as all available impressions are from the upper surface, and it is possible that the sporangia are distributed evenly or in irregular patches over the whole lamina. Individual sporangia are indistinct, they are about 0.05 mm in diameter. An annulus is faintly visible in some. Discussion: When Shirley erected this species, there were few others to compare it with. Subsequent authors, other than Jones & de Jersey (1947), have ignored it. No more recently erected species are synonymous with it although it is quite similar to D. tenutfoluum Stipanicic & Menendez, and D. ellenberg? Fabre & Greber, and less similar to D. davidtt Walkom and D. acutilobum (Braun) Schenk. All but the last are Gondwanan species, and it is felt that they form a natural, closely related group. Material Studied: Lectotype: GSQ F 166a (here designated), figured by Shirley Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 12 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE pore t» LD 7 — 3 SOO Fig. 3. 1-8 — Dictyophyllum bremerense Shirley, 1898 (1) GSQ F549, partly reconstructed x 34; (2) GSQ F12041 x %; (3) UQ F2685 x%; (4) UQ F64068 x14; (5) UQ F64068, sorus showing probable arrangement of sporangia x 634; (6) UQ F43868 x2; (7) UQ F64068 x 3%; (8) UQ F2668 x 3%. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST 13 (1898), pl. XIII, fig. 2; Denmark Hill, Ipswich, Qld; Late Triassic age. Paratype GSQ F 166, figured by Shirley (1898), pl. XIII, fig. 3; and counterpart UQ F 5981; from the same locality. Other Specimens: Denmark Hill, Blackstone Formation (upper part of the Ipswich Coal Measures) of Karnian age (de Jersey, 1972): GSQ F 549, 12033-12044; UQ F 2342, 2668, 2685, 5983, 5984, 44313; 5981 (counterpart of GSQ F 166) and 8199 both figured by Jones & de Jersey (1947; pl. 1 fig. 6; pl. iv, fig. 1b respectively) . Dinmore Quarry, Qld, Blackstone Formation: Mr N. Petty’s collection nos. 259, 264 a/b, 539, 607. Ipswich (details unknown) , Qld. SUP 20007 a,b. Dictyophyllum davidit Walkom, 1917 Fig. 1.7; Fig. 2.12; Fig. 5. 15-17 1917, Dictyophyllum davidt Walkom. Geol. Surv. Queensl., Publ. 257: 10, ple 3, figs 2 1924, Dictyophyllum rugosum Walkom. Mem. Queensl. Mus., 8:2, pl. 21, hige: lie 1954, Dictyophyllum bremerense, Shirley, in Derrington — unpublished thesis, University of Queensland: 45. 1975, Dictyophyllum davidat Walkom, zn Flint & Gould, J. Proc. R. Soc. INES. We 108) 71, pl. 1, fig. 3: Description: Frond palmate, with at least eight pinnae. The leaf is definitely petiolate, the petiole divides once into two main veins, which in turn divide over short intervals giving off four veins each to form the pinnae. The type-specimen has incomplete pinnae up to 50 mm long by 20 mm wide; in another big specimen they reach 100 mm long by 30 mm wide. Pinnae are only slightly pinnatifid, with small free “pinnules” only 4-5 mm long; it seems better to describe the pinnae as strongly dentate rather than pinnatifid, First order lateral veins arise at 70-75°, one for each “pinnule” or “tooth”, reaching their apex, slightly falcate. Veins of successive order cannot be clearly differentiated; they form a network of polygonal (4-7 sided) meshes. The meshes are somewhat elongated with their main axis parallel to the_pinna rachis, but become more isodiametric towards the margins and pinnae apices. The former are around 1.5 mm by 1 mm and the latter about 1 mm diameter. Only one fertile specimen is available; it shows elongated sori, 1.2-2 mm long by 0.8-1.2 mm wide, arranged mainly along the pinna rachis and first order lateral veins; very occasionally they seem to be on other parts of the lamina. Each sorus contains 25-30 sporangia but no details of these can be made out. Discussion: This species was believed to be closely related to the well known Dictyophyllum rugosum (L. & H.) and was sometimes mistaken with it. But the redescription given by Harris (1961) for this species shows very clearly that D. davidzz is a different species, with a general morphological similarity. Additionally the soral characters herein described for D. davidzt very strongly support this difference. As already stated it is felt that D. davidzz belongs to a natural “group” of species together with D. bremerense, D. tenuzfolium, D. ellenbergz and D. acutilobum. Material Studied: Holotype (here designated) : GSQ F 165, from “Challivet”, portion 28, parish Biarra, near Esk, Qid; Esk Formation of Middle Triassic age, largely Anisian (de Jersey, 1972). Other Specimens: Esk Formation of Middle Triassic age. “Challivet”, portion 28, parish Biarra, near Esk, Qld: GSQ F 168, 933, 12043; Various sites at or around Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 14 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE Fig. 4. 9-11 Hausmannia wilkinsit Walkom, 1928 (9) GSQ F1943a, combined with GSQ 8857 to give the reconstruction of a leaf 234; (10) GSQ F1943a, detail of venation x 234; (11) GSQ F8843, detail of venation X134. 12, 13 — Hausmannia (Protorhipis) sp. cf. H. (P.) deferrarisi Feruglio, 1937 (12) UQ F50306, detail of venation X 314 ; (13) UQ F50306, reconstruction of a leaf X 114. Wivenhoe Hill, parish Wivenhoe, Qld; UQ F 2010, 2046, 2051, 2359, 5812, 12853, 17068, 17069, 26754, 64177. Mandura 1:63 360 map sheet at grid reference 936629 (quoted from Derrington, 1954), Qld: UQ F 17394 a/b. Caboolture 1:63 360 map sheet at grid reference 645257, Qld: UQ F 17079 a/b. Portion 42, parish Wivenhoe, Qld: QM F 1468 (Walkom, 1924, pl. 21, fig. 1). Bundamba Group of latest Triassic to Middle Jurassic age (Cranfield & Schwarzbock, 1972), horizon within the Bundamba Group unknown. Precipice Creek (tributary of the Dawson River), 4 km SW of Rose’s Shack, Qld (Hill, in Shell Report, 1951) : UQ F 48643, 48654. Dictyophyllum acutilobum (Braun) Schenk 1917, D. acutzlobum (Braun) Schenk, in Arber, E.A.N., Paleont. Bull. N.Z. Geol. Surv. 6:34, pl. XII, figs 2-4. Discussion: I have not seen the original specimens described by Arber, but there is little doubt that they can confidently be regarded as belonging to the species. D. acutilobum has a rather wide distribution (Sweden, Germany, Persia and New Zealand) and was also quoted, without illustrations, by Zeiller (1875) from the “Rhaetic” of Chile; later Solms-Laubach, while describing plants from the same locality, quoted a different species of Dipteridaceae, but did not illustrate his species. The fragment found at Purga, Qld, described below is very similar to illustrations Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST 15 DS Jf Kae i i ~~ AC, ss 8. pe; vay X Fig. 5. 14, 18 — Thaumatopteris shirleyin.sp. (14) UQ F64280, reconstruction of part of the leaf based on the holotype x ¥ ; (18) UQ F64280, holotype X1. 15-17 — Dictyophyllum davidii Walkom, 1917 (15) GSQ F165, holotype x 234; (16) UQ F64177 x %; (17) GSQ F165, detail of venation of the holotype x 4. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 16 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE of this species figured by other authors, and therefore I consider that it most probably belongs here, but because of its fragmentary nature must be regarded as a comparison. Dictyophyllum sp. cf. D. acutelobum (Braun) Schenk Fig. 6. 16 Description: This pinna fragment is 28 mm long by 20 mm wide with two “pinnules” on each side. These are sub-triangular with a slightly curved midvein arising at about 45° from the rachis, and continuing to the apex of the “pinnules”. The free part of each “pinnule” is one third of the total width. From each midvein, secondary veins arise which, by successive divisions form a network of polygonal meshes. Those of the first order are about 4-5 mm diameter, and are filled with higher order meshes, the smallest one (highest order) are about 0.9- 1 mm diameter. Materzal Studied: Purga, Qld; Walloon Coal Measures, mainly Middle Jurassic: UQ F 5856. Thaumatopterts shirleyi n. sp. Fig. 2.9, 10; Fig. 5. 14, 18 Diagnosis: The biggest fragment represents a portion of pinna with a strong rachis 2 mm wide. First order lateral veins arise at 30° to enter the pinnules and reach their apex. The lamina forms a continuous wing along the rachis up to 22 mm wide, thereafter the pinnules are free. They are up to 72 mm long by 11 mm wide at the base, diminishing very slowly to 5 mm wide at the rounded apex. A small pinnule measures only 23 mm long by 5 mm at the base. Margins of the pinnules are crenated, each crena slightly asymmetrical, and about 3.5 mm long. Second order lateral veins (laterals of pinnules) emerge at 40-45°, and are spaced 4-6 mm apart. Those on the winged lamina are immediately dichotomized forming a net of polygonal first order meshes of 7 by 5 mm diameter, with their longest axis along the main rachis. Inside these, successively smaller meshes are formed, the smallest about 0.5 to 1 mm diameter; in many cases a free terminal veinlet in these ultimate meshes can be seen. In the pinnules only the small last order meshes stand out clearly. All specimens are sterile, but one of them shows some bulgings of the lamina which could correspond to underlying bodies (sporangia?). These bulgings coincide with the interior of the ultimate meshes. Discussion: 1 could not find a species among the Dipteridaceae which can be closely compared with T. shzrleyz. There are several forms with long “pinnules” but the lamina adjoining the rachis, the size and venation characters differ quite a bit. Although not known from complete leaves, T. shzrleyz seems to be one of those cases where it is difficult to decide between Thaumatopteris or Dictyophyllum as the best to house the species; the former name is preferred on account of the above- mentioned long “pinnules”’. Materzal Studied: Holotype: UQ F 64280, Paratypes: UQ F 64204 a/b from Dinmore Quarry, Ipswich, Qld in the Blackstone Formation (upper part of Ipswich Coal Measures) of mainly Karnian age (de Jersey, 1972). Other specimens: CTES-PB no. 2995, University of the Northeast, Corrientes, Argentina. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST 17 Fig. 6. 13, 14 — Hausmannia (Protorhipis) sp. cf. H. (P.) deferrariisz Feruglio, 1937 (13) UQ F50305 x234; (14) UQ F64201 434. 15 — Hausmannia wilkinsi Walkom, 1928 GSQ F1943a x2. 16 — Dictyophyllum sp. cf. D. acutilobum (Braun) Schenk, 1874 UQ F5868 234. 17 — Thaumatopteris shirleyi n. sp. UQ F64280, holotype x %. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 18 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE Hausmannia (Hausmannia ?) wilkinsia Walkom, 1928 Fig. 1. 1; Fig. 2. 11; Fig)6, Wb: Fig. 4. 910,11 1928, Hausmannia wilkinst Walkom. Proc. Linn. Soc. N.S.W., 53 (2): 148, pl. xiui, figs 3, 4. 1967, Hausmannia wilkinst Walkom, in White. Rep. Bur. Miner. Resour. Aust., 1967/78: fig. 3. 1972, Hausmannia wilkinsi Walkom, in White, Rep. Bur. Miner. Resour. Zlibyes MEP Me res (i. 7h Description: Complete(?) or half leaf(?) flabelliform, petiolate; the lamina is dissected into four main segments which in turn are again dissected but not so deeply. The four main segments correspond with each of the main veins into which the petiole splits; they dichotomize several times, each final vein reaching the apex of the ultimate portion of the dissected lamina. Veins of higher order arise at right angles and form a network of more or less quadrangular to 5-sided meshes of about 4-5 mm diameter. These are filled with successively smaller meshes, the smallest being about 1- 1.5 mm diameter. Discussion: The few more or less fragmentary specimens of H. wzlkinszz seem to show that it could be an intermediate form between the classical forms included in the subgenera Protorhipis and Hausmannia (s.s.) more probably inclined towards the latter. For its size and venation it was correctly placed in a separate species from those known to Walkom in 1928 and this difference still stands. Since erected by Walkom (1928), Hausmannia wilkinsa has only been used by White (see synonymy) for some fragmentary specimens, which appear to be correctly identified. These specimens came from the Nullawurt Sandstone Member of the Bungil Formation of Early Cretaceous age, and the Gilbert River Formation of Jurassic-Early Cretaceous age, in Queensland. Material Studied: Lectotype (here designated) GSQ F 1943, from Lower Camp, Plutoville, Cape York Peninsula, Qld, of Early Cretaceous age. Figured by Walkom Ce pl. xiii, fig. 4. Paratype GSQ F 1944, figured by Walkom (1928) pl. xiii, fig. One specimens: GSQ F 8843, 8844, 8851, 8857, 8858. All specimens come from the type locality. Hausmannia (Protorhipis) sp. cf. H. (P.) deferrarzsi. Feruglio, 1937 Fig. 1.6; Fig. 6. 13, 14; Fig. 4. 12, 13 1961, Hausmannia sp., in White, Rep. Bur. Mizner. Resour. Aust., 1961/146 fig. 15. 1966, Hausmannia (Protorhipis) sp., in Hill, Playford & Woods, Jurassic Fossils of Queensland (Queensl. Palaeontographical Soc.), pl. Jl, fig. 9. 1966, Haan sp., in White, Rep. Bur. Miner. Resour. Aust., 1966/111, fig. 1. 1974, Hausmanmaa sp. cf H. (Protorhipis) deferrariisi Feruglio, zn Gould, Proc. R. Soc. Queensl., 85 (3) : 35. Description: Leaf entire, composed of two half laminae separated by a deep lower and a short upper sinus. Each half lamina is more or less oval, 20-23 mm long by 15- 20 mm wide, margins markedly crenate. The strong petiole, which probably was Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST 19 originally at right angles to the lamina, gives off four primary veins into each half lamina. Each vein divides dichotomously several times to form a network of more or less quadrangular to hexagonal meshes of successively higher orders, the smallest measure about 0.4-0.5 mm diameter. The first dichotomy of the primary veins occurs halfway to the margin, and successive dichotomies along the primary veins branch at about 45°. Higher order veins and veinlets branch almost at right angles. All specimens are sterile. Discussion: Several good specimens of this species are in the University of Queensland collection, but even so it is difficult to establish definitely their complete identity with H. (P.) deferrarits Feruglio, which was originally described from the Middle to Upper Jurassic of Patagonia. There seem to be slight differences in size and venation characters, as well as considerable geographical separation between localities. As the Australian specimens are not completely identical with the Argentinian ones, the best procedure is to designate a comparison. Gould (1974) adopted a similar procedure. Some specimens quoted by White (1961, 1966) are too fragmentary to allow a definite determination, however they are most probably Early Cretaceous in age. The specimen originally figured by White (1966, fig. 1; reproduced here as Fig. 1. 6; the photograph was kindly supplied by Mrs White) seems to be fertile. It very strongly resembles H. (P.) papzlio (Feruglio) Herbst, a closely allied species. Material Studied: Walloon Coal Measures, mainly Middle Jurassic; Tannymorel Colliery, Tannymoreal, Qld. UQ F 50305, 50306, 64192, 64193, 64194 a/b, 64195- 64197, 64198 a/b, 64199, 64200. Mt Elliott Mine, Rosewood, Qld. UQ F 64201, 64202. Other Specimens: Walloon Coal Measures, mainly Middle Jurassic. Kleinton Clay Pit (via Toowoomba), Qld. QM F 2901. Near Kalbar, parish of Fassifern, Qld. QM F 2905. Hausmannia (Hausmannia) bulbaformis Douglas, 1973 1954, P?Angiosperm, in Medwell, Proc. R. Soc. Vict., 65: 21. 1969, Hausmannia sp. indet., in Douglas, Mem. geol. Surv. Vict., 28: 224; fig. 4, 2. 1969, Hausmannza sp., in Douglas, Mem. geol. Surv. Vict., 28: 232; pl. 42, fig. 4. 1973, Hausmannia bulbaformis Douglas, Mem. geol. Surv. Vict., 29: 96- Qs ok, Bei, sila, DS sales 7), Ce Discussion: Only two specimens are available and both are rather fragmentary. Specimen MUGD 3533A, the holotype, from the Koonwarra fish-beds shows some details of venation, thus it can be ascertained that the leaf belongs to the Dipteridaceae, and most probably to Hausmannza. It is a petiolate incomplete leaf, with several main veins dividing in a fan-like way which by successive divisions form a network of polygonal, slightly elongated, meshes about 1 mm in diameter. The lamina seems to have been quite thin and filmy. I think it is rather risky to erect a new species on such fragmentary material, but on the other hand it can be stated that the specimen does not resemble any of the known Australasian species of Hausmannia. Therefore I shall provisionally accept Douglas’ classification hoping that in the future more findings and better preserved material will justify the erection of this species. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 20 REVIEW OF THE AUSTRALIAN DIPTERIDACEAE Materzal Studied: Koonwarra Fish Beds, near Leongatha, Vic.; Wonthaggi 1:63 360 map sheet grid reference 3966, 2457. Early Cretaceous: MUDG 3533 A, B. Killara Bluff, allotment 4, section A, Parish of Killara, Vic. Early Cretaceous: MUDG 2014. cf. Clathropterts sp. Fig. 1. 4 Description: Fragments of lamina with typical “clathropteroid’” quadrangular network of meshes. None of the available specimens shows any part of the original margin, but the biggest one shows what appears to be the main rachis from which at least six first order (?) veins depart in a more or less palmate arrangement. They dichotomize at angles of 45°, and from their branches the veins of successive orders start at right angles to form the abovementioned quadrangular network with meshes about 3-3.5 mm diameter. Further away from the base, these meshes become more polygonal (5-6 sided), but remain about the same size. The ultimate meshes are about 1-1.5 mm each side, and again quadrangular at the base, and slightly more polygonal away from it. Materzal Studied: Cooroy 1:63 360 map sheet at grid reference 098146, Qld; Walloon Coal Measures, mainly Middle Jurassic: UQ F 23068-23072, UQ F 23074- 23076. Discussion: All known fragments are too small and fragmentary to allow a definite classification, but from the few diagnostic characters it can be concluded that the pieces most probably belong to Clathropteris. The venation is different from all other known species hitherto described for Australasia. ACKNOWLEDGEMENT j The John S. Guggenheim Foundation, New York, is thanked for award of a fellowship during tenure of which the present study was made. References ArseER, E. A. N., 1917.— The earlier Mesozoic Floras of New Zealand. Palaeont. Bull. N.Z. geol. Surv., 6: 1-80. CRANFIELD, L. C., and SCHWARZBOCK, H., 1972. — Nomenclature of some Mesozoic rocks in the Brisbane and Ipswich areas, Queensland. Queensl. govt Min. J., 73: 414-416. Dr JersEy, N. J., 1972.— Triassic Miospores from the Esk Beds. Geol. Surv. Queensl., Publ. 357, Pal. Pap. 32: 1-40. DERRINGTON, S. S., 1954.— The Geology of Murgon-Windera District. Unpubl. B.Sc. (Hons) Thesis, University of Queensland. Douc as, J. G., 1969. — The Mesozoic Floras of Victoria. Parts 1 and 2. Mem. geol. Surv. Vict., 28: 1-310. —— , 1973. — The Mesozoic Floras of Victoria. Part 3. Mem. geol. Surv. Vict., 29: 1-185. FABRE, J., and GrEBER, C., 1960.— Présence d’un Dictyophyllum dans la flore molteno du Basutoland (Afrique australe). Bull. Soc. géol. France, (7)2 (2) : 178-182. GouLp, R. E., 1974. — The fossil flora of the Walloon Coal Measures: a survey. Proc. R. Soc. Queensl., 85: 33-41. — , 1975. — The succession of Australian Pre-Tertiary megafossil floras. Bot. Rev., 41 (4) : 453-483. Harris, T. M., 1961.— The Yorkshire Jurassic Flora 1. Thallophyta — Pteridophyta. London: British Museum (Nat. Hist.). 212 pp. Hitt, D., 1951. — zm SHELL REPORT, q.v. —, PrayrorD, G., and Woops, J. T., 1966.— Jurassic Fossils of Queensland. Brisbane: Queensland Palaeontographical Society. 32 pp. Jones, O. A., 1948.— Triassic Plants from Cracow. Proc. R. Soc. Queensl., 49: 101-108. , and Dr Jersey, N. J., 1947.—The Flora of the Ipswich Coal Measures — morphology and floral succession. Pap. Dep. Geol. Univ. Queensl., (n.s.) 3(3) : 1-88. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 RAFAEL HERBST 21 SHELL REporT, 1951.— General Report on Investigations and Operations carried out by Shell Co. in the search for Ozl in Queensland. 1940-1951. Unpubl. rep. on file, Geol. Surv. Queensl. Library (CR 640). SHIRLEY, J., 1898. — Additions to the Fossil Flora of Queensland. Geol. Surv. Queensl., Publ. 128, Bull., 7: 1-25. Wa kom, A. B., 1917. — Mesozoic Floras of Queensland: The Flora of the Ipswich and Walloon Series (C) Filicales, etc. Part I. Continued. Geol. Surv. Queensl., Publ. 257: 1-67. ——, 1919.— Mesozoic Floras of Queensland, Parts 3 and 4. The Floras of the Burrum and Styx River Series. Geol. Surv. Queensl., Publ. 263: 1-77. ——, 1924. — On fossil plants from Bellevue, near Esk. Mem. Queensl. Mus., 8: 77-92. ——, 1928. — Fossil plants from Plutoville, Cape York Peninsula. Proc. Linn Soc. N.S.W., 53(2) : 145- 150. White, M. E., 1961.— Report on 1960 collections of Mesozoic plant fossils from the Northern Territory. Rec. Bur. Miner. Resour. Aust., 1961/146: 26 pp. (unpubl.). ——, 1966.— Report on 1965 plant fossil collections. Rec. Bur. Mzner. Resour. Aust., 1966/111: 10 pp (unpubl.). ——, 1967.— Report on 1966 collections of plant fossils from the Surat Basin, South-west Eromanga Basin, Delamere, Northern Territory: and Proserpine District of Queensland. Rec Bur. Miner. Resour. Aust., 1967/78: 17 pp. (unpubl.). —— , 1969. — Report on the 1968 collection of plant fossils from the Moolayember and Teviot Formations. Rec. Bur. Miner. Resour. Aust., 1969/51: 13 pp. (unpubl.). ——, 1972.— Mesozoic plant fossils from near Croydon, Queensland. Rec. Bur. Miner. Resour. Aust., 1972/31: 8 pp. (npubl.). ZEILLER, R., 1875. — Note sur les Plantes fossiles de La Ternera (Chili) . Bull. Soc. géol. France, (3)3: 572- 574. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 A new species of the Lztorza peronu Complex (Anura: Hylidae) from Eastern Australia A. A. MARTIN, G. F. WATSON, D. F. GARTSIDE, M. J. LITTLEJOHN and the late J. J. LOFTUS-HILLS MartTIN, A. A.. WaTSON, G. F., GARTSIDE, D. F., LITTLEJOHN, M. J., & Lorrus- HILts, J. J. A new species of the Lztorza peronzz complex (Anura: Hylidae) from eastern Australia. Proc. Linn. Soc. N.S.W. 103 (1), (1978) 1979: 23-35. A new species of hylid frog, Lztorza tylerz, is described from coastal eastern Australia. L. tylerz is a member of the L. peronz? complex; it differs from the other Australian members of the complex (L. peronzz and L. rothz) in adult morphology and mating-call structure. L. tyler? is sympatric with L. peronz and zn witro hybridization tests show the two species to be reciprocally genetically incompatible; they also differ biochemically. A. A. Martin, G. F. Watson, D. F. Gartside, M. J. Littlejohn and J. J. Loftus-Hills (deceased 11 June 1974), Department of Zoology, University of Melbourne, Parkuille, Australia 3052; manuscript received 5 July 1977, in revised form 15 November 1977. INTRODUCTION The Liztorza peroni group of species includes L. peronz in Australia, L. amboinensis, L. darlingtoni and L. everett2 in New Guinea and associated islands, and L. rothi in both Australia and New Guinea (Tyler, 1968; Menzies, 1976). In addition, Straughan (1966) and Tyler (1968) noted that an undescribed member of this complex occurs near Brisbane, Queensland. We have collected this undescribed species along eastern coastal Australia from south-eastern Queensland to Jervis Bay, New South Wales. The purpose of the present paper is to describe the new taxon and compare it with L. peronzi and L. rothz. Because we have encountered L. rothz only infrequently in the field, comparative data for this species include only morphology of preserved adults and larvae, and mating-call structure. For the new species and L. peroniz, however, we have considerable field and experimental data, encompassing adult morphology, mating- call structure, life history, larval morphology, reciprocal artificial hybridization tests, and electrophoresis of haemoglobins and plasma proteins. METHODS Adult Morphology Comparative morphological study was restricted to sexually mature males, and measurements were taken (to 0.1 mm) with vernier calipers or with a stereoscopic microscope and eyepiece micrometer. We have followed the methods and terminology of Tyler (1968), except that head length was measured to the mid-point of the tympanum. Abbreviations used in the text are: E-N/IN = eye to naris distance/internarial span; HL/HW = head length/head width; TL/S-V = tibia length/snout to vent length. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 24 A NEW SPECIES OF THE LITORIA PERONII COMPLEX Mating-Call Structure Mating calls were recorded in the field using a variety of tape recorders and microphones (e.g. EMI L2B, Nagra IIIBH, Tandberg 11-2 tape recorders; Beyer M- 69, Beyer M-88, Grampian DP-1 microphones). Where possible, the recorded specimens were collected and lodged in the Department of Zoology, University of Melbourne research collection. Wet-bulb air temperatures (which approximate those of small frogs calling from elevated positions) were taken at or near the calling sites. Recordings were analysed on an audiospectrograph (Kay 6061-B Sona-Graph) with playback on a Tandberg 11-2 tape recorder. One call of each individual was analysed, generally the last clear call in the recording sequence. Note and spectral characteristics were obtained from a note at or near the middle of each call. Characteristics of the calls were measured on the audiospectrograms using calibrated scales. Artificial Hybridization Tests and Life History Artificial hybridization tests were carried out in the field using the technique of Rugh (1962). In wtro crosses were made between individuals of the new taxon and L. peronzi, all collected from a sympatric breeding assemblage at Ryan’s Swamp, Caves Beach Reserve, A.C.T. For each interspecific cross a simultaneous intraspecific (control) cross was made. Progeny of control crosses provided material for life history descriptions; and for L. peronz additional larval material from Sarsfield, Victoria, derived from an zn wvo fertilization on 3rd December, 1965, was examined. Embryos and larvae were reared initially in the field under fluctuating temperature conditions and, on return to the laboratory, in Holtfreter’s Solution at 20 + 0.5°C. Larvae were fed on boiled lettuce. Larvae of L. rothz were collected at Kununurra, W.A. on 24th February, 1977. Methods of fixation, measurement and illustration of embryonic and larval material follow those of Martin and Littlejohn (1966). Blood Proteins Plasma proteins and haemoglobins from 12 individuals of L. peronz (6 from Caves Beach Reserve; 6 from Ourimbah, N.S.W.) and 9 of the new taxon (7 from Caves Beach Reserve; 2 from Ourimbah) were analysed. Animals were etherized and ventrally dissected to expose the cardiac cavity. A truncus arteriosus was cut, and blood collected in heparinized microhaematocrit tubes. The tubes were flame-sealed, centrifuged at low speed (approximately 25 g) for 10 minutes, then broken at the interface between plasma and cells. Electrophoresis was carried out on horizontal 5% acrylamide gel slabs using the method described by Gartside (1972). All separations were performed at room temperature using a continuous tris (tris-hydroxymethyl aminomethane) borate buffer (0.3 M tris; pH 8.7) and 250 volts constant voltage, provided by a Vokam stabilized power supply. Five yl of plasma were analysed from each individual. Electrophoresis of plasma was continued until bromphenol blue dye, added to human plasma controls on each gel, had migrated 7 cm from the origin: this took about two hours. Plasma proteins were stained in 1% amidoschwartz 10B (Chroma Gesellschaft) in 7% acetic acid. Haemoglobins, prepared by washing cells twice in 0.9% sodium chloride and haemolysing them in a mixture of equal parts of toluene and distilled water, were stained in o-dianisidine using the technique of Owen and Smith (1961). All blood samples were fresh when analysed. Proc. Linn. Soc. N.S.W., 108 (1), (1978) 1979 MARTIN, WATSON, GARTSIDE, LITTLEJOHN AND LOFTUS-HILLS 25 DESCRIPTION Litorza tylert n. sp. Types Holotype, mature male No. R64754, Australian Museum, Sydney; male paratypes: Nos R64755-R64764; female paratypes: Nos R64765, R64766. Type locality: Ryan’s Swamp, Caves Beach Reserve, A.C.T., 14 km S of Huskisson, Shire of Shoalhaven, N.S.W. (35° 09’ 45” S, 150° 40’ 00° E). Collected by D. F. Gartside, M. J. Littlejohn, J. J. Loftus-Hills, A. A. Martin, I. F. Spellerberg and G. F. Watson, 21st October, 1969. South Australian Museum Nos R12248, R12251, R12254, R13267A, R13267B, R13267C, R13338A, R13338B, R13338C, all mature males from the Ourimbah area, N.S.W., are also nominated as paratypes. Diagnosis The Lztorza peronz group is distinguished from all other frogs in Australia by the combination of the following characteristics : (1) well-developed webbing and discs on both fingers and toes; (ii) second finger longer than first ; (111) dorsum light grey to brown with minute green flecks in life; and (iv) posterior surfaces of thighs patterned with yellow and black. L. tylert can be distinguished from L. peronz and L. rothz by means of the following characters. In both L. peronz and L. roth there is a dark edging on the supratympanic ridge; this dark line is absent in L. tyler?. In L. rothi the axilla is black, and the posterior surface of the thigh is black with a few yellow spots. In L. peronz the axilla is yellow with large black spots, and the thigh is yellow with coarse 1. L. tyleri (10) 2. L.tyleri (9) 3. L. peronii (9) 4. L. peroni (9) 5. L. peronii (8) 6. L. peronii (10) 7. L. peroni (6) @ _L. rothi (10) 30 40 50 60 04 0.5 0.7 0.8 0.9 1.0 1.0 15 2.0 S-V (mm) TL/S-V HL/HW E-N/IN Fig. 1. Body dimensions and ratios of samples of Litorza tylert, L. peroni? and L. rotht. The number in parentheses after the species name is the sample size. The short vertical line is the sample mean and the horizontal line is the observed range of variation. The black bar represents the 95% confidence limits on either side of the mean and the open bar plus one-half of the black bar indicates one standard deviation of the mean. Sample localities are: 1 and 3, Caves Beach Reserve, A.C.T.; 2, Palm Grove, Ourimbah, N.S.W.; 4, Mallacoota, Vic.; 5, 5-19 km W of Coonabarabran, N.S.W.; 6, Narrandera, N.S.W.: 7, Yarrawonga area, Vic.; 8, Laura, Qld. Proc. Linn. Soc. N.S.W., 108 (1), (1978) 1979 26 A NEW SPECIES OF THE LITORIA PERONII COMPLEX black variegation. In L. tylerz the axilla is yellow with few or no black spots, and the thigh is yellow with fine black variegation. The texture of the back is rough and warty in L. peronz, much less warty in L. tyler?, and virtually smooth in L. rotht. The mating calls of the three species are distinctive. L. tylert has a greater S-V length than L. rotht (Fig. 1). Crosses between individuals of L. tyler: and L. peronit show the species to be reciprocally genetically incompatible (Table 1). TABLE 1 Results of zm wtro crosses between Litorza tylert and L. peroniz from Ryan’s Swamp, Caves Beach Reserve, A.C.T. Cross No. of Eggs No. Fertilized % Hatched* L. tyleri Q x L. peronii 8 Control 110 96 100 Experimental 67 67 0 Control 100 100 80 Experimental 61 61 0 L. peronii Q X L. tyleri 8 Control 40 40 70 Experimental 77 77 0 Control 65 65 60 Experimental 74 74 0 *Failure to hatch was associated with developmental breakdown, particularly abnormal neurulation. The three species may readily be identified by the use of the following key: 1. A black line bordering the supratympanic ridge; axilla black, or yellow with heavy black spots; posterior surface of thigh heavily marked. with blackoic, cos) ccc seve ecco: clot aaa aiecel tl mn Ue seep cee 2 Supratympanic ridge without a black line; axilla yellow, sometimes with 1-2 small black spots; posterior surface of thigh yellow withfine black. vanlegationtss 42 ..G) ai eine L. tylerz 2. Body length of mature males more than 42 mm; head length 76-88% of head width; back warty; posterior surface of thigh yellow withtheavy blackvarlegationien)...4c). 044 sci tae aee L. peroniz Body length of mature males less than 42 mm; head length 86-95% of head width; back smooth; posterior surface of thigh blackswithkartewayellow:Spotsiqcescci.) seas s.ct Gu-eencae Seen aaa en ie L. rothi Description A medium-sized species of moderately slender habitus (Fig. 2). Vomerine teeth between the internal nares; tongue broad and with a posterior nick. Head broader than long (HL/HW = 0.80-0.85) ; snout rounded. External nares much closer to tip of snout than to eye; internarial span less than distance between eye and naris (E-N/IN = 1.23 —1.59). Canthus rostralis slightly concave but not sharply defined ; loreal region sloping. Eye diameter slightly greater than distance from eye to naris, Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 MARTIN, WATSON, GARTSIDE, LITTLEJOHN AND LOFTUS-HILLS 27 Fig. 3. Palmar view of left hand and plantar view of left foot of holotype of Lztorza tylert. The bar represents 5 mm. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 28 A NEW SPECIES OF THE LITORIA PERONII COMPLEX and considerably greater than tympanum diameter. Tympanum prominent, bordered dorsally by a ridge running from behind the eye to the axilla. Webbing on fingers well-developed (Fig. 3); discs prominent, disc diameter being about 1.5-2 times phalanx width. Sequence of lengths 3 > 4 > 2 > 1. Hind limbs moderately short; TL/S-V = 0.43-0.49. Webbing extends to penultimate phalanx of fourth toe and almost to discs of other toes (Fig. 3) . Sequence of lengths 4>5 = 3 >2> 1. Inner metatarsal tubercle small, oval; no outer metatarsal tubercle. Dorsal skin mottled, light grey to medium brown, with minute emerald spots; texture finely warty. Ventral skin distinctly granular, off-white to yellowish. Gular region dusky brown in breeding males; no nuptial pads. Axilla with a yellow pterygial membrane, sometimes with 1-2 small black spots. Groin yellow with black mottling ; posterior surface of thigh with finely divided yellow and black markings. Dimensions of holotype are: snout-vent length, 43.4mm; tibia length, 20.3 mm; head length, 12.8 mm; head width, 15.2 mm; eye to naris distance, 4.3 mm; internarial span, 3.1mm; eye diameter, 4.7 mm; tympanum diameter, 3.3 mm.Dimensions and ratios of males in the type series are shown in Fig. 1. Dimensions of the two female paratypes are: snout-vent length, 47.6 and 45.6 mm; tibia length, 22.8 and 21.7.mm; head length, 14.3 and 14.1 mm; head width, 15.4 and 15.3 mm; eye to naris distance, 4.2 and 4.5 mm; internarial span, 3.7 and 3.7 mm; eye diameter, 4.9 and 4.4 mm; tympanum diameter, 3.8 and 3.5 mm. Variation and Comparison with Other Species The type series of L. tylert shows little morphological variation, and a sample of males from Ourimbah, N.S.W., conforms with the type series (Fig. 1). Also shown in Fig. 1 are selected dimensions and ratios of two samples of L. peronz from coastal N.S.W. and Victoria, three samples of L. peronz from inland N.S.W. and Victoria, and a sample of L. rothz from Laura, Queensland. L. rothz is the most distinctive member of the complex, having the smallest body size, longest legs and narrowest head. It also has the least warty dorsal skin and the greatest amount of black pigmentation in the axilla, groin and thighs. Males of L. tylerz are intermediate between those of L. rothz and L. peronz in body size and degree of wartiness of the dorsal skin. On the other hand, L. tylerz has the shortest legs and the least amount of black colouration in the axilla, groin and thigh region. Some differentiation between coastal and inland populations of L. peronzz is evident, with the coastal samples tending to have a greater body size, narrower head and higher E-N/IN ratio. Mating-Call Structure . The mating calls of the three species are of similar basic structure and consist of trains of regularly-repeated notes (Fig. 4), each of which is pulse modulated (Table 2) Although there are insufficient data to assess the effects of temperature on call structure in the L. peronz group, general trends are evident for vocalizations in other species of anurans (Littlejohn, 1978). There is an inverse relationship with temperature for durations of calls and notes, and a direct relationship for repetition rates of calls, notes and pulses. The numbers of notes per call and pulses per note, and dominant frequency, usually are not markedly affected by temperature. Hence, allowance must be made for temperature variation when comparing calls. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 MARTIN, WATSON, GARTSIDE, LITTLEJOHN AND LOFTUS-HILLS 29 t= me 60 Ty .€2329@ * * ~ = _ * — — kHz 0.2 04 0.6 0.8 10 1.2 14 1.6 18 20 seconds Fig. 4. Audiospectrograms of mating calls of: upper left, Lztorza tylerz, Palm Grove, Ourimbah, N.S.W.; upper right, L. rothz, Kununurra, W.A.; lower, L. peronzz, Palm Grove, Ourimbah, N.S.W. The trace for L. peroniz is the latter half of a call. Even so, it is clear that the calls of L. peronzz are much longer than those of the other two species, and that the dominant frequencies are slightly higher and note repetition rates slightly lower in calls of L. tyler? (Table 2). There are more notes in the call of L. peronzz, and more pulses in the notes of L. tyler (Table 2) . These latter meristic characters should be only minimally associated with temperature, and thus could be most useful in diagnosis. Life History and Larval Morphology The following descriptions are based on embryos and larvae reared from eggs of L.. tylert and L. peronz fertilized zn vitro at Caves Beach Reserve on 22nd October, 1969. The dimensions of seven newly-fertilized L. tyler? eggs (mean and range) are: embryo diameter, 1.32 mm (1.28-1.36); capsule diameter, 3.49 mm (3.20-3.60). The comparable figures for L. peroniz are 1.52 mm (1.48-1.56) and 3.00 mm (2.80-3.12). Thus L. tylert eggs are smaller, but with slightly larger capsules. In both species the animal pole is dark brown, the vegetal pole creamy-white, and the jelly capsule three-layered. When 44 hours old, embryos of both species were at stage 18 of Gosner (1960). Those of L. tyler? are slender, medium brown in colour, and approximately 3 mm in total length. The L. peronzz embryos are plumper, lighter in colour and shorter (length about 2.4 mm). The embryos of L. tylerz hatched after 57 hours, at stage 20, when their total length was about 4.8 mm. Three pairs of external gills are present. Newly-hatched embryos of L. peronz are very similar but slightly lighter in colour; they hatched at an age of 70 hours. All larvae were preserved at the age of 49 days, when they had reached stages 25-26. The dimensions (mean and range) of four L. tyler? larvae are: total length, 13.96 mm (12.80-14.72) ; tail length, 9.36 mm (8.80-9.76). Those of five L. peronz? larvae are: total length, 19.50 mm (17.00-21.90); tail length, 12.20 mm (10.90-13.90) . Apart from the difference in size, the larvae are generally similar, with those of L. tyler? having slightly heavier pigmentation and narrower tail fins. The mouth discs are not fully developed, but in the larvae of L. tyler? heavier pigmentation Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 A NEW SPECIES OF THE LITORIA PERONIJ COMPLEX 30 *s[[eo autos url yuasaid syed jenba omy , 0006-0481 O00F-955 IL-6 96-96 6 81-F LI S661 8LE 0°01 GS 8 LI 0016-0061 9S6-L96 O1-6 &6-G6 641-6 91 * 806 66 06 SLI 0006-0041 9L6-066 61-6 19-86 SS1-F Gl * 6&6 LOl &P 6 6L 0996-0516 LSS-S66 06-66 GL-19 66-68 * 6&6 L°66 89 0'6 0596-0586 666° LPS G6-S6 89-09 601-98 GLEG PLE 8&6 $9 £6 0S46-0966 89¢6-Lél O¢-LI 8éL-FLI VSD? * 98T 0°S6 66l LY (2H) (9as /sastnd) (0asut) (Das /sajou) Aouonbary ae y d1I0N /sasing uoneimng ney JURUTWIOG ‘day as[ng uonneday 210N 2[PPHN 129N 230N 2YjOL “JT pue wuosad “7 ‘21ajh} vI10T FO s[[ed SuNUL JO soTISI1gIOeIYS [eoIshyd IT20/S930N 06F-OL¢ . Ntew see Shh § 9°G6 VM ‘eiinununy 2YIOL VILOLT 0086-0616 Beas “M’S’N GGrG : aUeIs6 2 ‘yequiling ‘aa01n wyeg OLGS-OLES seh eroe ; : L¥9Z 6 GPI-So FL dA ‘quiog Asdi9g 2uoLad VIL0}T GP9-O1S “M'S'N ‘Inoqiepy G64 g G16 SJJOD FO MN WY 8°61 08-SSP : "M'S'N P19 Y 8°06 ‘yequiLing ‘aAo15 Weg SE§1e-OLel 3 ‘ TED) \/ OLSI Y O'eT Ol ‘Qarasay YIeag saarD 249)4} 1240]2T (Sasut) dZIS ys uoneinq adweg ‘dwoay i1y Aqipeoo7T pur satsads qInd 39M UALS JIv SasULI PUL SULIT Z ATGVL Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 MARTIN, WATSON, GARTSIDE, LITTLEJOHN AND LOFTUS-HILLS 31 Fig. 5. Stages in development of Lztorza peronz?, Sarsfield, Vic. A, stage 17; B, stage 20 (newly-hatched) ; Cand D, stage 29. The bar in each case represents 1 mm. Fig. 6. Mouth disc of larva at stage 29 of Litorta peroni?, Sarsfield, Vic. The bar represents 1 mm. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 32 A NEW SPECIES OF THE LITORIA PERONII COMPLEX of the labial papillae is evident. In both species the spiracle is sinistral and the anus dextral. Because a complete developmental series of L. tylerz from eggs to advanced larvae was not obtained, a series of L. peronz? stages from Sarsfield, Vic., has been used for illustration (Fig. 5). Available material of L. tylerz indicates that the embryonic and larval stages of the two species are similar, and Fig. 5 shows the general characteristics of both species. Live L. peronz larvae are pale golden-yellow, with a dark lateral stripe extending from the snout through the eye and along the dorsal edge of the body and tail musculature. The intestinal peritoneum is dark dorsally but silvery-white ventrally, and the oral and branchial areas are almost transparent but for scattered golden chromatophores. The most advanced larva in the series (stage 41) has a total length of 44.2 mm and tail length of 26.8 mm. The mouth disc (Fig. 6) is typical of most Australian hylid larvae (Martin and Watson, 1971). There are two rows of teeth in the upper labium and three in the lower; 2-3 rows of papillae extend around the sides and back of the mouth disc. The mouth formula is 1 1 1 1 9 1 The Sarsfield larvae took about 190 days to develop from fertilization to metamorphosis, but the larval life-span of L. tylerz is unknown. + A B Fig. 7. Two-banded haemoglobin phenotypes of A, Litorza peronzz and B, L. tyler?, both from Palm Grove, Ourimbah, N.S.W. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 MARTIN, WATSON, GARTSIDE, LITTLEJOHN AND LOFTUS-HILLS 33 Fourteen L. rothz larvae from Kununurra, W.A., range in total length from 26.5 mm (stage 25) to 67.7 mm (stage 41). In general morphology they closely resemble the larvae of L. peroni and L. tylert; however, their oral structure is distinctive. The lower jaw is much more robust than in the other species, and the third lower row of labial teeth is absent, or reduced to a small median structure bearing less than ten teeth. Artificial Hybridization Two zn witro crosses were made in both reciprocal combinations between L. tylerz and L. peroniz (Table 1). While the control crosses displayed some degree of abnormality, the experimental crosses in both combinations were characterized by COFFS @ HARBOUR oO oO TAREE @ © Call records @ Specimens examined 100 kilometres NOWRA @ pe Type Locality Fig. 8. The geographic distribution of Ltorza tylert. The inset shows the area of eastern Australia covered by the main map. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 34 A NEW SPECIES OF THE LITORIA PERONII COMPLEX total developmental breakdown. No hybrid embryo successfully completed neurulation. This high degree of incompatibility is suggestive of crosses between diploids and tetraploids (as in the American Hyla versicolor and H. chrysoscelis; Wasserman, 1970), and a study of karyotypes would be of interest. Blood Proteins Fig. 7 shows the typical electrophoretic pattern for haemoglobins, each species having two anodally-migrating bands. The mobility of the leading band is similar in both taxa, while the mobility of the second band is consistently different. Migration of the second band is much faster in L. tylerz than in L. peronzz. For plasma proteins, one set of bands appeared to be species-specific, although other bands in the complex plasma pattern differed in mobility or staining intensity between individuals. Despite this individual variability, bands which are presumed to be transferrins (based on experience with other hylids) are of similar mobility in every individual of both taxa. Breeding Biology Males of L. tylert have been heard calling in October, November and January. Males called from the banks of permanent ponds and swamps, or from elevated positions (0.5-1.0 m above the water) in emergent or marginal vegetation. Wet bulb air temperatures at calling sites ranged from 11.0-21.5°C. Reproductively active females have been collected in October. Distribution L. tylerz is distributed along the east coast of Australia from southern Queensland to the Jervis Bay area, N.S.W. (Fig. 8). The geographic range of L. tylerz is entirely included within that of L. peronz (see Moore, 1961, Fig. 50). Neither species is known to be sympatric with L. rothz, which is distributed along the northern and north-eastern coasts (Cogger, 1975). Etymology The species is named for Michael J. Tyler of the University of Adelaide, in recognition of his contributions to our knowledge of Australo-Papuan hylid frogs. CONCLUSIONS The three species of the Lztorza peronzz complex in Australia are readily distinguishable. The sympatric forms, L. tyler? and L. peronzz, are highly distinctive in most aspects studied, including mating-call structure, blood-protein characteristics, and genetic incompatibility. The apparently absolute level of post-mating isolation between them renders the occurrence of hybridization extremely improbable. In any case, hybrids should be recognizable on grounds of adult morphology (e.g. Martin, 1972), mating-call structure (e.g. Zweifel, 1968) or blood-protein patterns (e.g. Brown and Guttman, 1970). None of our data for these characteristics is suggestive of the occurrence of hybridization. ACKNOWLEDGEMENTS The bulk of travel and running costs were met by an ARGC Grant (No. 66/16172) to M.J.L., and by the University of Melbourne research vote to the Department of Zoology. The Ian Potter Foundation provided funds for A.A.M. to visit Kununurra. We thank the South Australian Museum for the loan of specimens of L. tylerz. Dr I. F. Spellerberg assisted in the field, and Ms M. Leahy prepared some of the figures. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 MARTIN, WATSON, GARTSIDE, LITTLEJOHN AND LOFTUS-HILLS 35 References Brown, L. E., and Gutrman, S. I., 1970.— Natural hybridization between the toads Bufo arenarum and Bufo spinulosus in Argentina. Amer. Midl. Nat., 83: 160-166. CocceErR, H. G., 1975. — Reptzles and amphibians of Australia. Sydney: A. H. and A. W. Reed. GarTSIDE, D. F., 1972.— The Litorza ewing? complex (Anura: Hylidae) in south-eastern Australia III. Blood protein variation across a narrow hybrid zone between L. ewzngz and L. paraewingz. Aust. J. Zool., 20: 435-443. GosneR, K. L., 1960.— A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16: 183-190. LITTLEJOHN, M. J., 1978.— Long-range acoustic communication in anurans: An integrated and evolutionary approach. Jn: Taylor, D. H. and Guttman, S. I. (eds), Reproductive Biology of Amphibians. New York: Plenum. Martin, A. A., 1972. — Studies in Australian Amphibia III. The Lzmnodynastes dorsalis complex (Anura: Leptodactylidae). Aust. J. Zool., 20: 165-211. , and LITTLEJOHN, M. J., 1966.— The breeding biology and larval development of Hyla jervisienszs (Anura: Hylidae). Proc. Linn, Soc. N.S.W., 91: 47-57. ——, and Watson, G. F., 1971.— Life history as an aid to generic delimitation in the family Hylidae. Copeza, 1971: 78-89. MENZIES, J. I., 1976. — Handbook of common New Guinea frogs. Wau: Wau Ecology Institute. Mookrg, J. A., 1961. — The frogs of eastern New South Wales. Bull. Amer. Mus. Nat. Hist., 121: 149-386. OwEn, J. A., and Smitu, H., 1961.— Detection of caeruloplasmin after zone electrophoresis. Clinica chim. Acta., 6: 441-444. Rucu, R., 1962. — Experzmental embryology. Minneapolis: Burgess Publishing Company. STRAUGHAN, I. R., 1966.— An analysis of species recognition and species isolation in certain Queensland frogs. Unpublished Ph.D. Thesis: University of Queensland. Ty er, M. J., 1968. — Papuan hylid frogs of the genus Hyla. Zool. Verhand., No. 96, pp. 1-203. WASSERMAN, A. O., 1970.— Polyploidy in the common tree toad Hyla verstcolor Le Conte. Science, 167: 385-386. ZWEIFEL, R. G., 1968.— Effects of temperature, body size and hybridization on mating calls of toads, Bufo a. americanus and Bufo woodhousez fowler. Copeza, 1968: 269-285. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 te a aad A New Species of Microscolex (Diplotrema) (Annelida: Oligochaeta) from New South Wales G. R. DYNE (Communicated by B. G. M. JAMIESON) Dyne, G. R. A new species of Microscolex (Diplotrema) (Annelida: Oligtochaeta) from New South Wales. Proc. Linn. Soc. N.S.W. 103 (1), (1978) 1979: 37-41. A new species of Microscolex (Diplotrema), the first record of the subgenus from New South Wales, is described and figured. This species, defining the known southerly limit of Dzplotrema in Australia, is shown to have close affinities with the subgeneric type, D. fragzlis. These species, together with an undescribed form from Ban Ban Springs, Queensland, appear to form a discrete species-group quite distinct from more northerly representatives of the subgenus. G. R. Dyne, Department of Zoology, University of Queensland, St. Lucia, Australia 4067; manuscript received 23 March 1976, in revised form 19 January 1978. INTRODUCTION The endemic Australian acanthodriline earthworm subgenus Microscolex (Diplotrema) contains sixteen described species, with a distribution extending from Gayndah (S.E. Queensland) to North-Western Australia, including a large nucleus of species in the Northern Territory. Spencer (1900) erected the genus Dzplotrema (type, D. fragilis) on the basis of the arrangement of the male terminalia, the openings of the vasa deferentia and prostatic ducts supposedly separate, but both on segment XVIII. Stephenson (1930) in devising an elaborate phylogenetic scheme, incorporated Diplotrema as a necessary intermediate between the primitive acanthodrilin condition (male pores on XVIII, two pairs of prostatic pores on XVII and XIX), and the more advanced megascolecin condition, with combined male and prostatic pores. However, re-examination of Spencer’s material (Jamieson, 1971) showed the original description to be grossly erroneous, D. fragzlis in fact having the acanthodrilin arrangement of male terminalia. The redefined Dzplotrema could not then maintain separate status from the type of Eodrilus, E. cornigravez Michaelsen, 1907, and consequently the large number of species formerly assigned to this genus were regrouped (Jamieson, 1971). Those forming an homogeneous assemblage with D. fragilis with respect to Australian endemicity and the possession of modified genital setae remained in Dzplotrema, whilst the majority of species were placed in Notzodrilus Michaelsen, 1950. Subsequent analysis (Jamieson, 1974) resulted in the relocation of the entire complex (as subgenera Notzodrilus and Diplotrema) in Microscolex Rosa, 1887, largely because the microscolecin arrangement of combined male and prostatic pores on XVII, characteristic of Mzcroscolex, had been shown in some instances to vary to the acanthodrilin condition intraspecifically (Pickford, 1932). A re-examination of the type-species of Notzodrilus, N. georgianus Michaelsen 1888, demonstrated sufficient differences from the type of Mzcroscolex, M. phosphoreus (Dugés 1837) to warrant subgeneric rank for Notzodrilus. This subgenus, within Mzcroscolex, embraced a small number of species with vesiculate Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 38 NEW SPECIES OF MICROSCOLEX (DIPLOTREMA) nephridia and a distinctive setal arrangement; Dzplotrema was incorporated as a third subgenus within Microscolex, differing from Notzodrilus primarily in the universal occurrence of genital setae within the species of Dzplotrema recognized at that time (Jamieson, 1974). Finally, a study of a number of new Dzplotrema species from the Northern Territory, Australia, which exhibited marked variation in the arrangement of genital setae, and which included species lacking such setae, led to a further redefinition of the subgenus (Jamieson and Dyne, 1976). Dzplotrema nevertheless retains a pronounced morphological and endemic identity. SYSTEMATIC DESCRIPTION Genus MICROSCOLEX Rosa, 1887 Subgenus DIPLOTREMA Spencer 1900 Emend. Diplotrema tyagarah sp. nov. Fig. 1 A-D, table 1 Material Examined: 149° 49’ E, 30° 07'S, 26 km north of Narrabri on Newell Highway, in black moist clayey soil under dry grass, W. Nash and R. Raven, 19 July 1975 — Holotype (H) (AM W 6622), paratype, Pl] (AM W 6623); 149° 37’ E, 29° 49’ S, 42 km south of Moree on Newell Hwy, in black clayey soil, just below surface in flat grassy country, W. Nash and R. Raven, 19 July 1975 — Paratype, P 2 (AM W 6624). 1 = 51, 49.5 + mm (posterior amputee); w (midclitellar) = 2.2, 2.1mm; s=142, 76+; (H, Pl). Form uniformly circular in cross-section; pigmentless buff in alcohol. Prostomium epilobous 4%, peristomium somewhat furrowed. First dorsal pore 7/8. Setae 8 per segment, in regular longitudinal rows throughout, setae a and b of XVII and XIX modified as penial setae; setae a and b modified as spermathecal genital setae in VII but absent from XVIII. Nephropores inconspicuous on the clitellum (in cd?) . Clitellum annular, strongly developed, XIII-X VII (extending dorsally into 4% XII) setae visible on the clitellum, intersegmental furrows faint, dorsal pores obscured. Male pores minute orifices in broad seminal grooves, in mid-XVIII, slightly later of b-lines; prostatic porophores 2 pairs, in XVII and XIX, on slight protruberances forming the four corners of a roughly square male field, delimited laterally by a pair of broad seminal grooves joining the prostatic porophores; within the male field a diamond arrangement of small elliptical markings, each a slightly glandular area with pore-like centre, in 17/18 and 18/19; a large transversely elliptical tumescence with raised rim and median ridge, with slightly sunken centre in XVI, pressing anteriorly to meet the indented posterior edge of the clitellum; a diffusely glandular area immediately posterior to the male field, extending to 20/21, and within it, a faintly visible marking, similar to those within the male field, median, in XIX, close to 19/20. Female pores conspicuous openings presetally in ab (closer to a) near intersegment 13/14, in XIV, surrounded by an elliptical, paler region on the clitellum. Spermathecal pores 2 pairs, closely paired in a-lines in 7/8 and 8/9, on protruberant lips. A large pair of glandular swellings in VII associated with the genital seta follicles, extending laterally to c, and filling the segment. Septa 6/9-10/11, slightly thickened, 6/7-8/9 moderately strongly thickened, 5/6 moderately thickened. Dorsal blood vessel single, continuous onto pharynx. Last hearts in XIII, supraoesophageal vessel IX-XIII, adherent to roof of oesophagus; commissurals in X-XIII larger than the remainder, and sending a narrow connective Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 G. R. DYNE 39 TABLE 1 Intersetal Distances in Segment XII of M. (Diplotrema) tyagarah % of circumference aa ab be cd dd dc cb ba u(in mm.) HOLOTYPE 9.3 2.1 14.5 2.1 51.8 1.9 16.0 2.3 5.25 PARATYPE 1 TES 22 14.3. 1.6 De 2a eles 15.0 2.0 5.51 PARATYPE 2 10.3. 2.5 15.3 2.0 51.3 2.0 14.0 2.5 4.37 mean 1O%5, 223 14.7. 1.9 51.4 1.9 15.0 2.3 5.04 1mm A 10 mu Fig. 1. M. (Diplotrema) tyagarah sp. nov. A.— Genital field of Holotype. B.— Right prostates of Holotype zn situ. C.— Right spermatheca (IX) of Holotype. D.— Mature genital seta from Holotype. Abbreviations used in illustrations and text: gl — glandular area; g.m — accessory genital marking; pr.p — prostatic pore; pr.d — prostatic duct; pr.g — prostate gland; p.s — penial seta; s — septum; s.tum — tumescence associated with genital setae; sem.gr — seminal groove; sp.amp — spermathecal ampulla; sp.d — spermathecal duct; sp.div — spermathecal diverticulum; sp.p — spermathecal pore; d — male pore; 2 — female pore; H — holotype; P — paratype; 1 — length; w — width; s — number of segments; u — total circumference. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 40 NEW SPECIES OF MICROSCOLEX (DIPLOTREMA) to both the dorsal and supraoesophageal vessels, remaining commissurals very small and dorso-ventral only. Gizzard small-medium, soft and compressible, in V; oesophagus VI-XIV, rather narrow, not well vascularized, with conspicuous rugae on its inner walls; intestinal origin in XV, with abrupt expansion, a definite typhlosole absent. Nephridia stomate holonephridia throughout, the medium = sized nephrostomes preseptal and usually in ab; the wide, thin-walled ducts entering the parietes slightly pre-setally in cd; the nephridial body invested in a high peritoneum appearing as a discrete sheaf of tissue in each segment; no tufting demonstrable in the pharyngeal region. Holandric; large flocculent sperm masses and very large, brightly iridescent sperm funnels in X and XI; seminal vesicles ill-defined, 2 pairs, one very small pair in IX, and a more prominent mass in XII, with apparent stalk-like connection to the funnels of the preceding segment. Coiled, tubular prostate glands 2 pairs, in XVII and XIX, somewhat flattened, and with stout muscular ducts; the anterior pair conspicuously the larger encroaching into XVIII. Penial seta follicles thin and transparent, with little copulatory musculature; a and 6 follicles conjoined ; the setae moderately long, ectally tapering shafts with some irregular blunt toothing on the distal eighth. Length of mature seta= 1.54 mm; midshaft diameter = 19.2 um (mean of 3). Ovaries in XIII, comprising small sheafs of oocytes and large pleated funnels. Spermathecae 2 pairs, subequal, in VIII and IX, each comprising a bulbous, ovoid ampulla joined by a short stalk to a clavate diverticulum, the walls of which are packed with innumerable iridescent spheroidal sperm chambers; whereas the ectal duct and diverticulum appear continuous, the stalked ampulla appears to be an appendage. Length of right spermatheca of IX (from apex of ampulla to pore) = 1.5 mm; total length = 5.6 x length of duct and 1.4 x length of diverticulum. Genital seta follicles conspicuous in VII, a and 6b follicles inseparable; the setae stout shafts conspicuously ornamented with deep longitudinal notching which becomes more elongate entally; length of mature seta = 0.65 mm, midshaft diameter = 22.2 um (mean of 2). Remarks: Diplotrema tyagarah is readily distinguishable from the remainder of the subgenus by the unique configuration of genital markings associated with the male field. The closest affinities of the species lie with the subgeneric type, D. fragzls, with which it shares such characters as similar general appearance of the genital field and clitellum, identical position of the first dorsal pore, lobular peritoneum-invested nephridial bodies, intestinal origin in XV, and a striking similarity in the form of the spermathecae. D. tyagarah is nevertheless separable on the basis of genital field details and setal ratio data. DISCUSSION Diplotrema tyagarah constitutes the first record of Diplotrema from New South Wales, extending the distributional range of the subgenus southwards by some 480 kilometres. The species thus forms the southernmost limit of a staggered chain of Diplotrema species extending from the Northern Territory, across to Cape York, and down the eastern coast of Queensland. Despite remoteness from northern relatives, D. tyagarah has maintained most of the characteristics which contribute to the homogeneity of the subgenus as a whole, including holandry, possession of peculiarly modified setae in the vicinity of the spermathecae, retention of penial setae, and consistency in the number of spermathecae and their diverticula. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 G. R. DYNE 41 Together with an undescribed species from Ban-Ban Springs (S.E. Qld), D. tyagarah and the subgeneric type, D. fragilis form a morphologically compact aggregate rather distinct from the more northerly representatives of the subgenus (wide Jamieson and Dyne, 1976 for descriptions and key). Radiation from a common ancestry is likely to have taken place within the last 10-50 thousand years, when increasing post-Pleistocene aridity was effectively isolating numerous earthworm populations. The members of species-groups have presumably resisted such isolating factors for a longer period, and remain closely related at the present time. More intensive collecting in and near the regions separating D. tyagarah from its close allies may reveal further isolates of similar affinities. ACKNOWLEDGEMENTS The author is indebted to Dr B. G. M. Jamieson, Department of Zoology, University of Queensland, for helpful advice and criticism during the course of this study. References Ducts, A., 1828. — Recherches sur la circulation, la respiration et la reproduction des annélides abranches. Annls Sci. nat. 15: 284-337. JaMiEson, B. G. M., 1971.— A review of the Megascolecoid Earthworm genera (Oligochaeta) of Australia Part 2 — The Subfamilies Ocnerodrilinae and Acanthodrilinae. Proc. R. Soc. Qd 82 (8) : 95-108. ——, 1974. — Generic type-species and other Megascolec¢dae (Annelida, Oligochaeta) in the Museum of Systematic Zoology, University of Turin. Boll. Mus. Zool. Univ. Torino 8: 57-88. ——, and Dyne, G. R., 1976.— The acanthodriline earthworm subgenus Microscolex (Diplotrema) in Northern Territory, Australia. Aust. J. Zool. 24: 445-476. MICHAELSEN, W., 1888. — Die Oligochaeten von Sud-Georgien. Jb. Hamb. wiss. Anst. 68: 55-73. PICKFORD, G. E., 1932. — Oligochaeta Part II. Earthworms. Dzscovery Reports 4: 265-290. Cambridge. Rosa, D., 1887.— Microscolex modestus, new genus and species. Boll. Mus. Zool. Anat. Comp. R. Univ. Torino 2, 19: 1-2. SPENCER, W. B., 1900. — Further descriptions of Australian earthworms, Part 1. Proc. R. Soc. Vict. 13 (n.s.), 1: 29-67. STEPHENSON, J., 1930. — The Oligochaeta. Oxford, Clarendon Press. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 hee righ = rns ae MT ee MCL SEN t era PR a at fy be hear he Re een ~. Me Sens The family Hypoderidae (Acari) in Australia ALEX FAIN and ROBERT DOMROW Fain, A., & Domrow, R. The family Hypoderidae (Acari) in Australia. Proc. Linn. Soc. N.S.W. 103 (1), (1978) 1979: 43-46. Mites of the family Hypoderidae are recorded from Australia for the first time: Neottialges (Pelecanectes) evansi Fain from Phalacrocorax carbo (Linnaeus) (Phalacrocoracidae) (Tasmania), N. (P.) tasmanienszs, n. sp., from P. fuscescens (Vieillot) and P. sulczrostris (Brandt) (Tasmania and New South Wales, respectively), and N. (P.) plegadzcola Fain from Threskiornis spinicollis (Jameson) (Threskiornithidae) (Victoria). A. Fain, Institut de Médecine Tropicale Prince Léopold, B-2000 Antwerp, Belgium and R. Domrow, Queensland Institute of Medical Research, Herston, Australia 4006; manuscript recetved 26 July 1977, in revised form 25 January 1978. Hypoderid mites are in general free-living, but the hypopial stage invades the subcutaneous tissues of birds and mammals (Fain and Laurence, 1974). A few species were listed from birds either of widespread distribution or introduced into Australia (Fain, 1967), but no specifically Australian material was previously known. We now record three species (of which one is new) taken from birds in south-eastern Australia. Genus NEOTTIALGES Fain Neottzalges Fain, 1966: 325. Type-species N. (N.) geopeliae Fain. Subgenus PELECANECTES Fain Pelecanectes Fain, 1966: 326. Type-species N. (P.) evans: Fain. Key to Australian species of Neottzalges 1. Genital sclerite either complete or represented only by ANCHO Mali SCAG A qnelyglON Gets tigen lees es ee eter 2 Genital sclerite interrupted at mid-length. Setae Cha SOOTY eis SOM caeis Ceci eye Ae Mee ees eerste tasmaniensis, N. sp. yee Gemealisclerte complete gira. ei. 2 a4 ne 5 evanst Fain Genital sclerite represented only by anterior half ..... plegadicola Fain Neottialges (Pelecanectes) evanst Fain N. (P.) evansi Fain, 1966: 327; 1967: 88; Cernj, 1969: 272; Pence, 1972: 435; Schwan and Sileo, 1978: 522. P. evanst (Fain) : Fain and Beaucournu, 1972: 374. Material. Many hypopial specimens from “subcutaneous nodule”, black cormorant, Phalacrocorax carbo (Linnaeus) (Pelecaniformes: Phalacrocoracidae), near Scottsdale, Tasmania ili. 1977, R. W. Mason. In authors’ institutes. Notes. The only previous records were from various cormorants, Phalacrocorax spp., in England, France, Kenya, Cuba and the U.S.A. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 44 THE FAMILY HYPODERIDAE (ACARI) Re wu! {QO Figs 1-2. Neottialges (Pelecanectes) tasmanienszs holotype hypopus. 1. Venter. 2. Dorsum. Neottzalges (Pelecanectes) tasmaniensts, n. sp. (Figs 1-5) Materzal. Holotype and 20 paratype hypopial specimens from “subcutaneous fascia infiltrated by mites”, white-breasted cormorant, P. fuscescens (Vieillot), Beechford, Tasmania, 10.vi.1973, B. L. Munday. Holotype and one paratype (on same slide, holotype @ g., i.e. to left, nearer red label) in Australian National Insect Collection, CSIRO, Canberra; remaining paratypes in authors’ institutes. Many specimens (not types) , same data, in spirit. Also many hypopial specimens (not types) found “subcutaneously on legs, neck and in particularly large concentration on the breast”, little black cormorant, P. sulctrostris (Brandt), Lake Cowal, New South Wales, 18.v.1977, W. J. M. Vestjens. In ANIC and authors’ institutes. Hypopus. Idiosoma not constricted at mid-length, 630 x 270 um (holotype), 640 X 265 um (paratype). Cuticle poorly sclerotized; dorsum and coxae without punctate shields. Palposomal sclerite wider (33 wm) than long (15 um) (Fig. 1). Epimera I fused; sternum proper forked posteriorly, 36 wm long. Genital suckers longer than wide, not divergent posteriorly. Genital sclerite represented by anterior T- Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 ALEX FAIN AND ROBERT DOMROW 45 shaped and posterior inverted T-shaped parts. Dorsal setae d,-3, ; and /, very long (175-225 um); J,-; and d, short (not above 25 ym) (Fig. 2). Legs not unduly shortened. Tarsi I-IV 39, 42, 72 and 60 um long, respectively. Tarsus I with eight piliform setae, three spines and two solenidia (terminal spine on tarsi I-II shorter than corresponding segment) (Fig. 3); tarsus III with eight piliform setae and one terminal spine (Fig. 4) ; tarsus IV shorter than tarsus III, with three spines (two basals at same level) and one elongate, barbulate terminal seta (Fig. 5). Notes. The new species is known only in the hypopial stage. It belongs to a small group characterized by the centrally interrupted genital sclerite and the short to very short setae /,_, and (usually) d,. The group previously contained three species: N. (P.) bassani (Montagu) from a sulid (Europe and southern Atlantic), and N. (P.) ajajae Fain and N. (P.) montagui Fain, both from a threskiornithid (transported from North America to the Antwerp Zoo) . The new species differs from N. (P.) bassanz in having seta d, set more laterally, tarsi III-IV of unequal lengths and no spine on tarsus III at midlength. It differs from N. (P.) ajajae in having seta d, much shorter, tarsi III-IV of more unequal lengths and a different disposition of spines on tarsus IV. Nearest to N. (P.) montaguz, to which it runs in Fain’s key (1967), it differs therefrom in having the body smaller, the palposomal sclerite much wider than long, the sternum longer, tarsi I-II much shorter (39 and 42 um vs 48 and 51 um, respectively) , a different disposition of basal spines on tarsus IV (at same vs different levels) and the trochanteral setae much longer. Neottzalges (Pelecanectes) plegadicola Fain N. (N.) plegadicola Fain, 1966: 326. N. (N., sic — recte P.) plegadicola Fain: Fain, 1967: 94. Figs 3-5. Neottialges (Pelecanectes) tasmanienszs hypopus. 3. Leg I. 4. Leg III. 5. Leg IV. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 46 THE FAMILY HYPODERIDAE (ACARI) Materzal. Hypopial specimens from “subcutaneous tissue around the cloaca . . . in clumps and in fairly large numbers”, straw-necked ibis (adult), Threskzornis spinicollis (Jameson) (Ciconiiformes: Threskiornithidae), rookery on Lake Corangamite, Victoria, ix.1973, J. H. Arundel and K. Harrigan. In authors’ institutes. Notes. The only previous record was from the glossy ibis, Plegadzs falcinellus (Linnaeus) (Threskiornithidae), in Belgium. This bird species also occurs in Australia. References Cerny, V., 1969. — The hypopi of Hypoderidae (Acarina: Sarcoptiformes) parasitizing Cuban birds. Folza parasit., Praha, 16: 271-274. Fain, A., 1966.— Note sur les acariens nidicoles 4 deutonymphe parasite tissulaire des oiseaux (Hypodectidae: Sarcoptiformes) (note preliminaire) . Revue Zool. Bot. afr., 74: 324-330. ——, 1967.— Les hypopes parasites des tissus cellulaires des oiseaux (Hypodectidae: Sarcoptiformes) . Bull. Inst. r. Sct. nat. Belg., 43 (4) : 1-139. ——, and BEAUCOURNU, J., 1972.— Observations sur le cycle €volutif de Pelecanectes evansi Fain et description d’une espece nouvelle du genre Phalacrodectes Fain (Hypoderidae: Sarcoptiformes) . Acarologia, 13: 374-382. ——, and LauRENCE, B. R., 1974.— A guide to the heteromorphic deutonymphs or hypopi (Acarina: Hypoderidae) living under the skin of birds, with the description of Ibzszdectes debilis gen. and sp. nov. from the scarlet ibis. J. nat. Hist., 8: 223-230. PENCE, D. B., 1972.— The hypopi (Acarina: Sarcoptiformes: Hypoderidae) from the subcutaneous tissues of birds in Louisiana. J. med. Ent., 9: 435-438. ScHWAN, T. G., and SiLEo, L., 1978.—Neottzalges (Pelecanectes) evanst (Sarcoptiformes: Hypoderidae) parasitizing a white-necked cormorant in Kenya. J. med. Ent., 14: 522. Corrigendum : These Proceedings, vol. 101, page 200, line 38 — delete “(?)” Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 An Early Triassic Megafossil Flora from the Lorne Basin, New South Wales W.B. K. HOLMES and S. R. ASH Homes, W. B. K., & Asn, S. R. An Early Triassic megafossil flora from the Lorne Basin, New South Wales. Proc. Linn. Soc. N.S.W. 103 (1), (1978) 1979: 47- 70. A small late Early Triassic flora is described in this paper from the lower part of the Camden Haven Group in the Lorne Basin of eastern New South Wales. It includes 17 identifiable species and several unidentified forms. New taxa in the flora are Coniopteris? ramosa sp. nov., Cladophlebis carne sp. nov., Dicrozdium voiseyi sp. nov., the supposed micro- and megasporophylls of Dicrocdzum zuberi (Szajnocha) Archangelsky var. feistmanteli (Johnston) Retallack, Karzbacarpon feistmantelz sp. nov., and Pteruchus barrealensis (Frenguelli) var. fezstmanteli comb. et var. nov., and the coniferous cone Voltziopsts townrovi sp. nov. The flora is similar and probably contemporaneous with that from the upper part of the Narrabeen Group of the Sydney Basin of New South Wales. W. B. K. Holmes, “Noonee-Nyrang”, Gulgong Road, Wellington, Australia 2820 (Hon. Research Fellow, Department of Geology, University of New England, Armidale, Australia 2351) and S. R. Ash, Department of Geology-Geography, Weber State College, Ogden, Utah, U.S.A. 85508; manuscript received 12 May 1978. INTRODUCTION Plant fossils of Early Triassic age occur at several localities in the Lorne Basin of northeastern New South Wales (Fig. 1). A few authors (Carne 1897, 1898; Voisey 1939; Pratt 1970) noted these occurrences, some listed the fossils they found but none of the fossils has been described. In this report we describe the recognizable plant megafossils that occur in the Lorne Basin at five localities near Laurieton (Fig. 1) and correlate the flora with the better known Triassic floras of the adjacent Sydney and Clarence-Moreton Basins of New South Wales. The fossils described are in the collections of the Geology Department, University of New England, Armidale (UNEF) , The Mining and Geological Museum, Sydney (MMF), Australian Museum, Sydney (AMF), and the Geology and Geophysics Department of Sydney University (SUGD). PREVIOUS INVESTIGATIONS The geology of what is now known as the Lorne Basin was first studied in 1896 by J. E. Carne, a Geological Surveyor with the Geological Survey of New South Wales. During the course of his investigation Carne discovered plant megafossils in a bed of grey shale at the base of the coastal headland called Camden Head (Fig. 1) in rocks now believed to be of late Early Triassic age and referred to the Camden Head Claystone (Fig. 2). Carne collected a few fossils and these were examined by W. S. Dun, Assistant Palaeontologist and Librarian of the N.S.W. Geological Survey, who reported (in Carne 1897, 1898) that the collection contained: Thznnfeldza odontopteroides (Morris) Feistmantel 1878, Alethopteris lindleyana Royle 1833, Equisetum, cycad frond — probably Ptilophyllum, Gleichenites?, Cardiocarpum Brongniart 1828, and Phyllotheca sp. He also reported that the collection contained a poorly preserved fern which was somewhat similar to Dédymosorus (Gleichenites) Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 48 AN EARLY TRIASSIC MEGAFOSSIL FLORA \e Clarence : Port X Scat : Macquarie ‘Bosin’,) New ; 0 England wy ss 0 5 lO km A vA | = Fold Belt S 1696 ES Port Lorne & Mecatenel Dyers Head Gorden Haven t Basin 1694-5 roup es 2 71693 age es ie @) (Laimisiony Perpendicular ault cs Sydney ee ea Fines ees 6 Basin =o x eS Camden Head UNE Fossil locality &Q SYONEY E oe fe) sli 200 km AUSTRALIA Eoney Fig. 1. Map showing the location of the Lorne Basin and major tectonic features in southeastern Australia and a more detailed locality map. The tectonic map is adapted from the Tectonic Map of Australia (Geol. Soc. Australia, 1971) and the locality map is adapted from the Hastings 1:250 000 geological sheet (1966) . 2 © 52 oS 5 Scale 0 Oo NG) 5m Ss ® HIS 2 D\C| o (dp) D> ie Legend Q)Y)s — El SI) 3 1695 FIT] 5 ; ® UNE Fossil sE ‘ 7) 6 55 locality @o m|S| § Es SI5|§ = O Grey Red Claystone Claystone ? Weaker Conglomerate Fig. 2. Stratigraphic section of the lower part of the Camden Haven Group exposed near the base of Grants Head, N.S.W. showing the positions of localities 1694 and 1695. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 W.B.K. HOLMES ANDS. R. ASH 49 gletchenozdes Oldham and Morris 1862. Carne concluded that the fossils showed that the strata could be correlated with the Clarence Coal Measures of Triassic age. During the 1930s A. H. Voisey studied the geology of the Lorne Basin and proposed (1939) that name for the area. Voisey (1939) also proposed the name Camden Haven Series for the Triassic rocks in the basin and collected plant fossils from what was apparently Carne’s locality at the base of Camden Head. Some of them were exhibited at the Ordinary Monthly Meeting of the Linnean Society of New South Wales on 29th September 1937, according to the Proceedings of the Society for that year (p. xlii). Voisey’s collection was studied by A. B. Walkom who reported (in Voisey 1939) that it contained Thinnfeldza ferstmantela Johnston 1895, Cladophlebzs sp. and a seed. Walkom also suggested that the fossil determined to be Thinnfeldia odontopterozdes by Dun (in Carne 1897, 1898) was probably the same as the fossil he called T. fezstmanteliz. In the same report Voisey mentioned the occurrence of Triassic plant fossils at several additional localities in the Lorne Basin but he did not identify any of them. We have been unable to locate the collections of Carne and Voisey. Goodwin (1962) studied the geology of part of the Lorne Basin and reported the occurrence of plants at several localities including one on the north side of Perpendicular Point (probably near locality 1693 of this paper). She, however, did not describe or illustrate any of the fossils. Packham (1969) summarized the geology of the Lorne Basin and named the Triassic strata there the Camden Haven Group. G. W. Pratt, who has studied the geology of the Lorne Basin for many years collected a few plants from Carne’s locality at the base of Camden Head. They were identified by Dr John Pickett (in Pratt 1970) as Cladophlebis australes (Morris) Seward 1904, Dzcrozdzum fezstmantelz (Johnston) Gothan 1912, ?>Willzamsonza sp.., and ?Neocalamites. Pratt and Herbert (1973) discussed the geology of the Lorne Basin in some detail and subdivided the Camden Haven Group into three formations: Camden Head Claystone (at the base), Laurieton Conglomerate, and Grants Head Formation (at the top). They suggested that the sequence was of Early Triassic age and correlated it with the upper part of the Narrabeen Group in the Sydney Basin. Helby (1972) has studied the pollen and spore content of sediments from two localities in the Lorne Basin. A sample from UNE locality 1583 yielded only three forms, none of which was of significance for dating purposes and a sample from the Grants Head locality 1695 yielded 10 species of palynomorphs. Helby (1973) stated that the association resembled the Aratrisporites tenuzspinosus Playford 1965 assemblage‘of late Early Triassic age in the Sydney Basin. G. J. Retallack has collected from the Camden Head Claystone at the base of Camden Head and has examined several collections of plant megafossils made by others from this locality. He lists (Retallack 1977) the following species: Cladophlebis sp., Dicrotdtum lancifoloum var. lineatum (Tenison-Woods) Retallack 1977, D. zubert (Szajnocha) Archangelsky 1968, “Pterorrachis” barrealensis Frenguelli 1942, Umkomasia sp., Lepidopteris madagascariensis Carpentier 1935, and Voltzzopszs wolganensis Townrow 1967. In the same report Retallack places this flora in the Dicroidetum zuberi Association. Bocking (1977) collected specimens from the Grants Head localities. He identified several species of Dzcrozdium from fragmentary remains. The best preserved specimen is included below as D. dubzum var. australe (Jacob & Jacob) Retallack. STRATIGRAPHY OF THE CAMDEN HAVEN GROUP The lowest member of the Camden Haven Group, the Camden Head Claystone, Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 50 AN EARLY TRIASSIC MEGAFOSSIL FLORA consists principally of red-brown claystone and siltstone and small amounts of grey fossiliferous claystone, sandstone and conglomerate. It is thick bedded and ranges from 0 to at least 75 m in thickness (Pratt and Herbert 1973, p. 108). The Camden Head Claystone which is thickest along the coastline thins westward and is absent in the western part of the Lorne Basin. The Camden Head Claystone unconformably overlies Palaeozoic rocks and is overlain by and intertongues with the Laurieton Conglomerate. Bocking (1977) regards the Camden Head Claystone as being wholly contained within the Laurieton Conglomerate. The Laurieton Conglomerate is massive bedded and yellow to brown in colour. It is composed of rounded to subangular pebbles and cobbles in a sandy to clayey matrix and ranges from 45 m to 210 m in thickness. The unit is thinnest along the coast and thickens to the west where it overlaps the underlying Camden Head Claystone and rests unconformably on Palaeozoic rocks in the western parts of the Lorne Basin. The Grants Head Formation, the uppermost member of the Camden Haven Group, consists of greyish sandsone, conglomerate and fossiliferous siltstone. It is thin to medium bedded and may be in excess of 150 metres thick. LOCALITIES The fossils described here were collected from five localities in the Camden Haven Group near Laurieton on the north coast of New South Wales, as shown on Fig. 1. Each locality is described below together with the assigned University of New England locality number. The grid reference numbers are taken from the Camden Haven 1: 100 000 sheet. 1583 Camden Head Claystone. In a lens of hard grey siltsone at the base of Camden Head. This is the locality from which Carne (1897, 1898), Voisey (1939), Pratt (1970) and Retallack (1977) have collected. It is commonly called the Perpendicular Point locality but the original locality descriptions given by these authors indicate that their collections actually came from the base of Camden Head not Perpendicular Point. G. R. 846984. 1693 Camden Head Claystone. In thin bedded grey siltstone exposed at the base of the sea cliffs on the north side of Perpendicular Point. G. R. 850990. 1694 Camden Head Claystone. In a bed of nodular grey siltstone near the base of the sea cliffs at Grants Head. About 19 m below the Laurieton Conglomerate of the Camden Haven Group (Fig. 2). G.R. 854034. 1695 Camden Head Claystone. In a lens of nodular grey siltstone in the sea cliffs at Grants Head. About 10 m below the Laurieton Conglomerate of the Camden Haven Group (Fig. 2). G.R. 854034. 1696 Grants Head Formation. In thin-bedded siltstone in the sea cliffs at the northern end of Bartletts Beach. G.R. 850045. SYSTEMATIC PALAEOBOTANY Division Tracheophyta Class Lycopsida Order Lepidodendrales Genus Skzllzostrobus Ash 1979 Skilliostrobus australis Ash 1979 Fig. 3, 1 and 11 Description. This recently described cone is represented in the flora by isolated examples of its distinctive sporophylls. The proximal portions of the sporophyll are Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 W.B.K. HOLMES ANDS. R. ASH 51 wedge shaped and have a maximum width of 12 mm and a length of about 1 cm. The limbs have a width of about 3 mm and a length of about 4.5 mm but the apices of the limbs are missing. Megaspores characteristic of S. australas were extracted from one of the megasporophylls (Fig. 3,11). Discussion. S. australis occurs in the lower part of the Gosford Formation of Early Triassic age near Terrigal, New South Wales and in the Parmenteer Supergroup near Hobart, Tasmania (Ash 1979). Material. AMF 59457 from locality 1695 and specimen UNEF 15510 from locality 1583. Lycopod Stems of Uncertain Position ?Lycopod stem a Fig. 3,2 Description. A stem 12 cm in length and 1.5 cm in width shows a surface covered with oval scars approximately 2.5mm X 1 mm. The preservation of this fossil makes it difficult to identify. It does superficially resemble the stem of Pleuromeza longicaulis (Burges) Retallack (1975, fig. 5A) so we presume that it is a lycopod stem. The scars, however, are somewhat similar to those that occur in Psaronzus and it is possible that the fossil is a fern stem. Material. Specimens AMF 59416 and UNEF 15509 from locality 1583. Lycopod stem b Fig. 3,3 Description. A fragment of a lepidendroid stem which has three vertical rows of slightly concave rhomboidal leaf cushions in a steep spiral arrangement. The cushions are attenuated vertically, the height being about 16 mm and the width 8 mm. The top of each cushion is level with the midpoint of the horizontally adjacent cushions. Structureless carbonaceous material crusts the surface of each cushion and no leaf scars or pits are preserved. Discussion. The orientation of the specimen in life is unknown and the description given above applies to the specimen as aligned in the illustration. Lepidodendroid stems with similar but smaller rhombic leaf cushions also occur in the Newport Formation at UNEL 1436 specimen UNEF 13829, and in the Basin Creek Formation at UNE locality 1489 (WBKH collection) . Material. AMF 59418 from locality 1583. Division Sphenopsida Equisetaceous stems Fig. 3,4 Description. Fragments of longitudinally ribbed equisetaceous stems have been collected from all recorded localities. At locality 1693 these fragments constitute the bulk of recognizable remains. Carne (1897) recorded Equzsetum and Phyllotheca sp. and Pratt and Herbert (1970) listed ?Neocalamites from locality 1583. None of the specimens examined by us have any diagnostic characters preserved that would allow a specific determination. Stems vary in width from about 2.5 mm to 30 mm and the number of ribs varies from 8 to 12 per cm. On most stems the internodes are apparently very long as few nodes are present. On stems with nodes the longitudinal ribs continue without interruption through the node. Rounded scars on these nodes may be branch or leaf bases. These stems could be referred to Paracalamites Zalessky 1927. Discussion. Similar stems have been recorded as Phyllotheca austials Brongniart, Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 52 AN EARLY TRIASSIC MEGAFOSSIL FLORA 1 Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 W.B.K. HOLMES ANDS. R. ASH 53 Schizoneura sp. or Neocalamites sp. from the Esk Beds (Hill et al. 1965. pl. T1, fig. 5; Walkom 1924, pl. 16, fig. 1), the Narrabeen Group (Walkom 1925, pl. 24, fig. 1) and Leigh Creek (Chapman and Cookson 1926, pl. 19, figs 2-6 and pl. 20, fig. 1). Materzal. MMF 16038, AMF 59424 and UNEF 15511, UNEF 15513, UNEF 15515 from locality 1583. Nodal Diaphragms Fig. 3,5 Two specimens are in the present collection. One is damaged and has some vascular bundles 1 mm apart still attached. The external diameter is approximately 18 mm. The other specimen consists of a circular sediment-filled cavity 10 mm in diameter surrounded by a 6 mm wide ring of about 28 segments. The specimen illustrated here in Fig. 3 compares rather closely with the nodal diaphragm from the Late Triassic of Greenland which Harris (1931, pl. 3, fig. 13) has called Equzsetites doratodon, and those illustrated as Neocalamites cf. carreri from the Middle Triassic Blackstone Formation of Queensland in Hill et al. (1965, pl. T1, 18K 8) Materzal. AMF 59423 and UNEF 15512 from locality 1583. Class Filicopsida Order Filicales Family Dicksoniaceae Genus Conzopteris Brongniart 1849 Coniopteris ?ramosa sp. nov. Fig. 4,1 and2 Diagnosis. Bi-(?tri-) pinnate leaf; penultimate segments pinnate-pinnatifid. Pinnae opposite, acutely triangular. Pinnules sub-opposite, oval, entire; imbricate near rachis but coalescing distally; a single vein entering each pinnule and forking several times. Description. Portion of a bi- (?tri-) pinnate leaf 30 mm in length with base and apex missing. Six pairs of opposite, acutely-triangular pinnae about 12 mm in length are attached almost at right angles to this length of rachis which is 1 mm in width. The pinnules near the main rachis are oval and contracted basally; 1.3 mm wide and 1.7 mm long; inserted at an angle of approximately 60°; margins entire and overlapping. The pinnae taper evenly throughout their length so that each succeeding pair of sub- opposite pinnules is slightly smaller and in the distal half become coalescent. A single vein enters the pinnule and branches up to seven times, radiating throughout the lamina. The specific epithet ramosa alludes to this character. Holotype. AMF 59428 and its counterpart AMF 59429. Locality. 1695 in grey siltstone lens in sea cliffs at Grants Head, N.S.W. Horizon. Camden Head Claystone, Camden Haven Group, Late Early Triassic. Discussion. Vhis fragment of a sterile fern leaf resembles in outline, sterile leaves of some other Triassic Conzopteris species, e.g. C. walkomi Frenguelli (1950, fig. 4) from Argentina and C. lobata (Oldham) Walkom (1925, pl. 29, figs. 4-6) from the Fig. 3. Note: Specimens natural size unless stated otherwise. Those photographed under kerosene are marked with an asterisk (*) . 1 Skzlliostrobus australis Ash, *, Detached megasporophyll, AMF59457. 2 ?Lycopod stem a, *, AMF 59416. 3 Lycopod stem b, AMF 59418. 4 Equisetaceous stem, AMF 59424. 5 Nodal diaphragm, AMF 59423. 6 Microphyllopteris sp., AMF 59417. 7 Cone?, AMF 59420. 8 Circular ribbed object, AMF 59419, x2. 9 Dispersed seed c, AMF 59421, <2. 10 Dispersed seed b, *, AMF 59422, X2. 11 Side of a megaspore ex- tracted from the detached megasporophyll attributed to Skzlliostrobus australis Ashin 1 AMF 59457, x 100. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 54 AN EARLY TRIASSIC MEGAFOSSIL FLORA ry : BR, ~ PF ¢ oY Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 W.B.K. HOLMES ANDS. R. ASH 55 Sydney Basin, but these species differ by having pinnular venation which branches once only. Sterile leaves of the Jurassic C. hymenophyllozdes (Brongn.) Seward have pinnules with veins that branch several times but the pinnules are lobed or finely divided (Harris 1961, fig. 53). Fertile specimens of C. ? ramosa are required to establish the correct generic assignment of this leaf. 2Coniopteris burejensis (Zalessky) Seward 1912 Fig. 5, 1 and2 Description. A sterile fern pinna, 28 mm in length and tapering only slightly. Pinnules alternate, rhomboidal with acute apices 2 mm wide and 3 mm long; inserted on rachis at an angle of about 45°; acroscopic lobe larger than the basiscopic lobe. Pinnule midrib giving off a pair of opposite or three or more alternate lateral veins which sometimes branch again before reaching the margin. Discussion. This specimen agrees in shape and venation with sterile fragments of C. burejensis from Yorkshire illustrated by Harris (1961, figs. 51 F-H) and is doubtfully assigned to that species. C. burejenszs is widespread in the Jurassic of Eastern Asia, Siberia and Europe. Material. AMF 59435 from locality 1583. Fern Form Genera of Uncertain Position Genus Cladophlebis Brongn. emend. Frenguelli 1947 Cladophlebis cf. mendozaensis (Geinitz) Frenguelli 1947 Fig. 4, 3 Description. A poorly preserved fragment of one side of a fern pinna 20 mm in length with parts of five oblong-lanceolate pinnules attached by the whole of their bases to the rachis, has venation in the pinnules somewhat characteristic of this species from the Late Triassic of Argentina (Frenguelli 1947, pl. 11, fig. 3). The majority of secondary veins visible on this specimen leave the pinnule midrib at an angle of approximately 45° and divide into two branches; the upper branch then divides again. Materzal. AMF 59426 from locality 1583. Cladophlebis carnez sp. nov. Fig. 4,4 and5 Diagnosis. Bipinnate leaf; pinnae opposite, linear. Pinnules catadromic; the first pinnule attached in the angle between the main rachis and the pinna rachis; the rest of the pinnules opposite or sub-opposite. Pinnules of irregular length, oblong or slightly tapering, obtuse; entire; inserted on pinna rachis at 75°-90°; midrib decurrent, persisting to or almost to the apex. Lateral veins catadromic, alternate to sub-opposite, mostly once divided. The basal lateral veins on either side of the midrib differ in shape from the other lateral veins. The first basiscopic vein is inserted at right angles to the base of the decurrent midvein or directly to the pinna rachis; dividing once and passing straight or slightly recurved to the margin. The first acroscopic lateral vein is attached close and parallel to the pinna rachis, then decurving to follow parallel around the acroscopic basal lobe of the pinnule. This vein is simple, or if Fig. 4. Note: Specimens natural size unless stated otherwise. Those photographed under kerosene are marked with an asterisk (*). 1-2 Coniopteris ? ramosa sp. nov., *, AMF 59428, holotype, 1, X 1.2, x3. 3 Cladophlebis cf. mendozeansts (Geintz) Frenguelli, *, AMF 59426, x3. 4-5 Cladophlebis carnez sp. nov., *, AMF 59425, holotype, 4, *1.5, X2. 6 Cladophlebis sp. indet., *, UNEF 15501. 7 Cladophlebis sp. indet., *, counterpart of speci- men in 6, AMF 59427, x2. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 AN EARLY TRIASSIC MEGAFOSSIL FLORA 56 Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 W. B. K. HOLMES ANDS. R. ASH 57 forked, the two branches run close and parallel. The following lateral veins leave the midrib at 20°-30°, dividing once near the midrib into two branches which diverge only slightly and curve to the margin at 45°-60°. The last two pairs of apical lateral veins are usually undivided. Description. Bipinnate leaf with portions of four opposite pinnae preserved. The pairs of pinnae are inserted at 75° and spaced about 23 mm apart on the main rachis which is 2.5 mm in width. The maximum preserved length of a pinna with a distal portion missing is 6 cm. Pinnules catadromic; first pinnule attached in the angle between the main and pinna rachis; following pinnules opposite, becoming subopposite to alternate distally. Pinnules slightly variable in length and width, 10 mm-13 mm X 3.5 mm-5 mm; oblong or slightly tapering; obtuse, entire; attached to the pinna rachis at 75°-90°. Pinnules decurrent, acroscopically contracted; the decurrent base of the next pinnule almost, or slightly, coalescing with the previous pinnule. Pinnule midrib decurrent and persistent almost to the apex. Lateral veins catadromic, alternate to sub-opposite; the first basiscopic and first acroscopic lateral veins differ in their course from the following veins. The first basiscopic lateral vein is inserted at right angles to the base of the decurrent midrib or directly to the pinna rachis; dividing once and passing straight or slightly recurved to the margin. The first acroscopic lateral vein is attached close and parallel to the pinna rachis; and then decurving to follow parallel around the acroscopic basal lobe of the pinnule; simple, or if forked, the two branches run close and parallel. The following lateral veins are inserted at 20°-30°; dividing once near the midrib; the branches diverging only slightly and curving to meet the margin at 45°-60°. The last one or two apical pairs of veins are usually undivided. 18- 21 lateral veins leave the midrib, with 34-40 reaching the margin. This species is named in honour of J. E. Carne [1855-1922] who made the first recorded collection of fossil plants from the Lorne Basin. Holotype. AMF 59425. Locality. 1583 in grey siltstone lens at base of sea cliffs on southern side of Camden Head. Horizon. Camden Head Claystone, Camden Head Group. Late Early Triassic. Discussion. C. carnez bears a superficial resemblance to Merzanopteris major Feist. 1881, and to Cladophlebis roylet (= Alethopteris lindleyana Royle) (Arber 1905, figs 34 and 32 respectively) and may be the fern leaf collected by Carne and identified by Dun as the latter species (Carne 1897) . Our species differs from both the above species by having twice as many lateral veins which divide only once — commonly twice in C. roylez. In Mertanopteris major the pinnules are coalescent, with the lower lateral veins anastomosing between pinnules. The C. roylez of Jack and Etheridge (1892, pl. 17, figs 3 and 4) and Merizanopteris major of Tenison-Woods (1883, pl. 8, figs 2 and 3) both differ from the types of these species and also from Cladophlebis carnez. The fragment referred to C. sp. cf. oblonga Halle, 1913a in Bourke et al. (1977, fig. 3.2) is C. carnez. C. oblonga differs in having undifferentiated lateral veins at the base of the pinnules. Cladophlebis sp. indet. Fig. 4,6 and7 Fig. 5. Note: Specimens natural size unless stated otherwise. Those photographed under kerosene are marked with an asterisk (*) . 1-2 Coniopteris burejensis (Zalessky) Seward ?, *, AMF 59435, 1, x1. 2, X2.5. 3-4 Dicroidium zuberz (Szajnocha) Archangelsky var. feistmanteli (Johnston) Retallack. 3, AMF 59430, *, 0.5.4, AMF 59431. 5 D. dubium var. australe (Jacob & Jacob) Retallack. Apical portion of leaf, *, AMF 59432. 6-7 Lepidop- teris madagascariensis Carpentier. 6, AMF 59433, X0.5. 7, *, AMF 59434. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 58 AN EARLY TRIASSIC MEGAFOSSIL FLORA Description. Some fragments of linear fern pinnae have very closely spaced, ovate to slightly falcate alternate pinnules with entire margins and rounded apex. The pinnules are closely spaced but not conjoined and are attached by the whole of the base and are slightly decurrent on the pinna rachis; 10-14 mm long and 4-6 mm wide; inserted at an angle of 50°-60°. Midrib slightly decurrent then curving outwards and continuing straight to within 1 mm of the pinnule apex where it divides into 2 or 3 secondary veins. The lateral veins are arranged alternately; the first basiscopic vein emerging at right angles from the decurrent base of the midrib or even lower from the rachis itself; dividing once and decurving through 70°-90° to the pinnule margin. The rest of the lateral veins, on both sides of the midrib, depart from it at an angle of about 30° and curve outwards to meet the margin at angles of 50°-75°. In the proximal two- thirds of the pinnule the lateral veins dichotomize twice in the first half of their length, but approaching the apex there is only one dichotomy or even undivided veins. About 20 lateral veins leave the midrib, dividing into 50 or more around the margin. Material. AMF 59427 and UNEF 15501 from locality 1583. Discussion. This pinna fragment is similar in dimensions to C. carnez sp. nov. but differs in having falcate pinnules; more divisions in the lateral veins and in the first acroscopic lateral vein being similar to the other lateral veins. The pinnules are shorter and broader and more closely spaced than those of C. australis (Morris) Halle 1913b, C. oblonga Halle has similarly shaped and spaced pinnules but differs in the lateral venation. In C. oblonga the lateral veins normally fork once only about midway from the midrib to the margin. Genus Microphyllopterts Arber 1917 Microphyllopteris sp. Fig. 3,6 Description. A fragment of a bipinnate leaf with portions of four parallel and closely spaced linear pinnae is placed in this form genus which was created by Arber for leaves formerly placed in the Glezchenztes genus but without evidence of fructifications to support their affinities with the present genus Glezchenza. The longest pinna fragment is 5.5 cm in length and decreases in width from 6 mm to 4 mm. The pinna rachis tapers gradually from 1 mm in width. The closely spaced opposite pinnules are 2 mm wide and 2.5 mm long and are attached by the whole of their base; slightly falcate to triangular with an acute apex near the primary rachis but decreasing gradually in size and becoming more rounded distally. No venation is preserved. Discussion. This leaf may be the same species as Carne’s specimens which were identified by Dun (Carne 1897, 1898) as Glezchenites or Didymosorus (Gleichenites) glecchenotdes Oldham and Morris 1862. The apices of the pinnules on our specimen are not rounded as in specimens of Glezchenites gletchenoides from the Rajmahal Series of India (Oldham and Morris 1862, pl. 25 and pl. 26, figs 1 and 3), or as narrow and acute as Microphyllopteris acuta Walkom 1919, from the Burrum Series of Queensland; Herbst (1974, p. 80) suggests that M. acuta may be a conifer. Our fossil compares closely with Todites narrabeenensis Burges (1935) from the Narrabeen Group of New South Wales and also in outline with the fertile fern Gleichenites wivenhoensis Herbst 1974 from the Esk Formation of Queensland. Material. AMF 59417 from locality 1583. Class Cycadopsida Order Pteridospermales Family Corystospermaceae Genus Dicrozditum Gothan, emend. Townrow 1957 Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 W. B. K. HOLMES ANDS. R. ASH 59 Dicroidium vorseyt sp. nov. Fig. 7,1 and2 Diagnosis. Pinnate frond with linear pinnules confluent on rachis. Lateral veins in pinnae well spaced, leaving midrib at acute angle and dichotomizing once, rarely twice. Description. Small pinnate forked(?) frond. Portion preserved (base and apex missing) 70 mm long and 40 mm wide. Main rachis 1.5 mm wide and gradually tapering upwards. Pinnae alternate, leaving rachis at angle of 40°-50°, bases confluent giving rachis a winged appearance; linear, with straight or slightly undulate margins, parallel for three-quarters of length then slowly tapering to a bluntly acute apex. Pinnae in mid-portion of frond 2.5-4 mm in width and 30-40 mm in length. A straight midrib runs the whole length of each pinna, the first lateral vein attached directly to the main rachis, the rest of the lateral veins leaving the midrib at an angle of 30°-45° and soon dichotomizing once with a very acute angle and continuing straight to the margin; occasionally a vein branches a second time; about 6 pairs of veins leave the midrib in 10 mm. This species is named in honour of Professor A. H. Voisey who pioneered the geology of the Lorne Basin. Holotype. AMF 59447 and counterpart AMF 56478. Paratype. AMF 59448. Locality. 1583 in grey siltstone lens at base of cliffs on southern side of Camden Head, N.S.W. Horizon. Camden Head Claystone, Camden Haven Group. Late Early Triassic. Discussion. Retallack (1977, microfiche Frame G7) identified specimen AMF 56478, the counterpart of the holotype of D. vozseyz as D. lancifolium var. lineatum (Tenison- Woods) Retallack. However, the lectotype of D. lancifolium var. lineatum, SUGD SUM 34 which was illustrated by Tenison-Woods (1883, Pl. 3, fig. 6) is a poorly preserved lower portion of a forked leaf which differs from D. vozseyz in having apparently grouped and numerous veins in acutely-triangular pinnules with contracted bases. D. pinnis-distantibus (Kurtz) Frenguelli 1943 appears to be coriaceous in contrast to the thin textured pinnae of D. vozseyz. The former species also has more distantly spaced and more tapering pinnae with a different venation. The sparse venation of D. vozseyz resembles that of Alethopterts medlicottzana Oldham (in Feistmantel 1876), from the Jabalpur Group of the Upper Gondwanas of India (Feistmantel 1886, pl. 1, figs 12-14). In outline D. vozseyz resembles Supaza linaerifolia White 1929, fig. 1, from the Permian Hermit Shale of Arizona. Thinnfeldia indica var. aquilina and T. indica var. falcata of Shirley (1898, pl. 6, fig. 2 and pl. 7, fig. 2) are close to Decrozdium votsey: but differ in the size and shape of the pinnae and in the venation. Walkom (1917a, p. 24) took Shirley’s illustrated specimen of Thznnfeldza indica var. falcata as the type for T. acuta — later Dicroidium acutum (Walkom) Frenguelli 1943. Retallack (1977, p. 271) synonymized D. acutum under D. lancifoltum var. lineatum (Tenison-Woods) Retallack. We consider D. acutum to be significantly different and should retain its specific status. Dicrotdium zubert (Szajnocha) Archangelsky, var. fezstmanteli (johnston) Retallack 1977 Fig. 5,3 and4 Description. Bipinnate leaves with forked rachis. Although none of the specimens is complete the fragments suggest that the larger leaves were at least as wide as 20 cm and 35 cm or more in length. The primary rachis of the leaf is 4-12 mm in width and has a punctate or papillate surface. The pinnae range from 2 cm to 10 cm in length Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 60 AN EARLY TRIASSIC MEGAFOSSIL FLORA and from 8 mm to 20 mm in width; attached to the primary rachis at an angle of 45°- 80°. The pinnae are divided to the pinna rachis into somewhat elongated subtriangular or rhombic pinnules 3 mm to 6 mm wide and 4 mm to 12 mm long, except towards their distal ends and near the leaf apex where they are coalescent and resemble similar portions of leaves of D. dubzwm and D. lanczfolcum. The pinnules are inserted at an angle of about 45° and have subacute to rounded or somewhat flattened apices. Several veins enter each pinnule from the pinna rachis and follow a curving path to the pinnule margins. Each vein typically forks twice before reaching the margin. Discussion. This species is a common element in the flora and is marked by a great variation in the size of the leaves. These fossils compare more closely with the variety D. zubert var. feistmanteliz (Johnston) Retallack 1977, but in some specimens the pinnules are more rhombic than rounded and come close to D. zubert var. zubert (Szajnocha) Archangelsky 1968. However, both the supposed male and female reproductive organs of the var. fezstmanteliz differ significantly from the South American fructifications attributed to the var. zwberz, which suggests that all our leaves should be placed in the var. fezstmantelzz. D. zuberi var. fezstmantela also occurs in the Narrabeen Group in the Sydney Basin (Walkom 1925, 1932), the Basin Creek Formation (Retallack et al. 1977) and the Esk Beds of Queensland (Walkom 1924). Material. AMF 59430, AMF 59431, and UNEF 15514 from locality 1583. Dicrotdium dubium var. australe (Jacob & Jacob) Retallack 1977 Fig. 5,5 Description. A medium sized bipinnatifid leaf. Pinna bases decurrent, pinna lobes rounded, venation indistinct. Discussion. Fragments of bipinnatifid pinnae are common at locality 1695 and probably belong to this species. The apical pinnae of the bipinnate D. zwberz group of leaves become bipinnatifid and may be confused with the above species. Materzal. AMF 59432 from locality 1583 and SUGD 81.8 from locality 1695. Genus Karibacarpon Lacey 1976 Type species: Karzbacarpon problematicum Lacey 1976 Emended Diagnosis. Megasporophyll composed of a stout rachis and several oppositely arranged lateral and a pair of terminal seed-bearing branches. Seeds contained in stalked, dorsiventrally flattened, fan-shaped, ribbed cupules. Seeds oval, large, ridged, with acute bifid micropyle. Empty cupules opening to form large star-shaped structures composed of 5 to 9 lobes. Karibacarpon feistmanteli sp. nov. Fig. 6, 1-8 Diagnosis. Small Karibacarpon fructification on a bipinnate megasporophyll. Closed cupules with inconspicuous ribs, 5-9 in number, usually 6. Mature cupules with lobed or petaloid margins, each enclosing one ovate platyspermic ovule. Empty cupules opening to form star-shaped “flowers”. Description. The megasporophyll consists of a stout primary rachis with opposite lateral branches bearing two or more pairs of opposite pedicellate cupules and a terminal pair of single cupules. Secondary rachis 3 mm or more in width and up to 5 cm in length. Pedicels of varying length and 3 mm to 4 mm in width at point of attachment to side of cupule which is of a very thick woody (?) texture. Closed cupules circular, oval or reniform, with 5 to 9, usually 6 inconspicuous ribs radiating from the Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 W.B.K. HOLMES ANDS. R. ASH 61 pedicel across the upper surface. The dorsal surface averages 18 mm X 14 mm and the sides are 2 mm deep (as compressed). Maturing cupules develop lobed margins with the ribs as suture lines between the lobes. The lobes become increasingly separated and form elongate acute “bracts” clasping the ovule. The lobes of empty cupules are expanded and when compressed form star-like “flowers”, 2 cm to 4 cm in diameter. The 5 to 9, usually 6 petal-like lobes are more or less acutely pointed and range from 6 mm to 16 mm in length, with a maximum width of 3.5 mm at the base. Most expanded cupules have been converted to a thick mass of structureless granular coal, suggesting that the organ was highly lignified. Occasionally the surface of the coal has a peculiar puckered appearance with irregular rounded ridges and furrows crossing the lobes at right angles. Walkom’s specimen (1925, p. 29, fig. 9) and Fig. 6, 6 of this paper illustrate examples of this type of preservation. The mature ovules are ovate and vary only slightly from the average size of 12 mm x 8 mm. A low medial ridge or fold extends the length of most ovules. The linear curved micropyle 1 mm in length is rarely preserved. Syntypes. AMF 59436 portion of megasporophyll (counterpart UNEF 15521) AMF 59439 closed cupule AMF 59437 lobed cupules with seeds (counterpart AMF 59438) AMF 59440 expanded cupule AMF 59445 dispersed seed Type locality. 1583 in a grey siltstone lens at the base of sea cliffs on southern side of Camden Head, N.S.W. Horizon. Camden Head Claystone, Camden Haven Group. Late Early Triassic. Discussion. The genus Karibacarpon was proposed by Lacey (1974) and erected (Lacey 1976) for a large cupulate fructification found in association with dispersed seeds and a large leafed Dzcrozdtum, D. narrabeenense var. bursellit (Lacey) Retallack 1977. Lacey (1976, p. 10) believed they were all derived from the one plant. Fertile organs of a similar nature have also been recorded by Lacey (1976, p. 11) in association with D. lancifolium. Both forms, which occur in the Molteno Succession in Rhodesia, are larger than Kar¢bacarpon feistmanteli sp. nov., have more ribs and are relatively much deeper, although the latter feature may be a result of preservation due to different types of enclosing sediments (Walton 1936). Thomas (1933) described three genera of female cupulate fructifications; Umkomasia, Pilophorosperma and Spermatocodon, from the Molteno Beds of Natal. On the evidence of similar cuticles and on the fact of association, he included these reproductive organs and the leaves of Dicrorditum Gothan, Xylopteris Frenguelli, Pachypterzs in part, and Johnstonza Walkom, in the family Corystospermaceae. In Umkomasia and Pilophorosperma the arrangement of the fertile branches is similar to K. feistmantelz sp. nov., but in those two genera the cupules are much smaller and divide into only two lobes by clefts in the plane of branching. The margins of older cupules of Spermatocodon have 3-4 small lobes but the infloresence is spirally branching. The largest cupule of S. sewardz Thomas (1933, fig. 31b) is less than one- quarter the average size of the cupules of K. fezstmantelzz. Frenguelli (1944) attributed cupulate ovuliferous fructifications to Dzcrozdzwm zubert (=Zuberia zuberz) and compared them with Lagenostoma. These cupules were borne on slender branches and opened into three lobes, (see Frenguelli 1944, figs. 10, 11) giving them an appearance quite different from Karibacarpon fetstmantel sp. nov. Examples of expanded cupules have been long known in the Early Triassic floras of the Sydney Basin. From the Newport Formation, Walkom (1925, Pl. 29, figs 7-9) illustrated specimens which he referred to ?Wzllm to 7 mm long and | mm or less wide, has the general appearance of V. angusta (Vv. ‘“om) Townrow (1967, fig. 1b), but our leaves are twice the size of that species. Material. AMF 59452 from locality 1695. Order Ginkgoales Form Genus Rhipidopsis Schmalhausen 1879 Rhipidopsis 2narrabeenensis Walkom 1925 Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 AN EARLY TRIASSIC MEGAFOSSIL FLORA ge i o> ¢ & i > oe % . ~ Soe a> Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 W.B.K. HOLMES ANDS. R. ASH 67 Fig. 7,8 Description. An apical portion of a wedge-shaped leaf, divided vertically into two segments, resembles an illustrated specimen from the Newport Formation of the Sydney Basin (Walkom 1925, pl. 30, fig. 4). The segments are 43 mm and 30 mm in width across the truncate apex; the length is estimated to have been from 8 to 10 cm. Straight veins about 1 mm apart radiate to the apex. There is no evidence of anastomosing to suggest that these leaves may be Chzropteris Kurr, but they may be better placed in Gznkgophytopsis Boureau. Burges (1935, p. 262, fig. 9) illustrated more complete specimens, probably from the same locality as Walkom’s. Other similar leaves are Rhipidopsis densinervis Feist. from Argentina (Kurtz 1921, Pl. 6, figs 51 and 53) and possibly R. genkgozdes Schmalhausen (?) var. sussmilchz Dun (1910, Pl. 51) from the roof shales of the Bulli Coal Seam under Sydney. Material. AMF 59455 from locality 1583. Plantae Sedis Incertae Stem? Fig. 7, 10 A poorly preserved example of a ? stem with scars possibly representing leaf bases resembles a specimen from the Narrabeen Group figured by Walkom (1925, Pl. 30, fig. 2) as ?Williamsonza sp. Materzal. AMF 59456 from UNEL 1583. Cone? Fig. 3,7 An ovate, woody organ 52 mm X 38 mm suggests a coniferous female cone similar to Penus or Sequoza. Oblong — spathulate bracts or scales 10 mm X 5mm are attached at an angle of 60° to a central axis. Materzal. AMF 59420 from UNEL 1583. Circular Ribbed Object Fig. 3,8 A round object 8.5 mm in diameter with a depressed centre and about 10 raised spoke-like ridges radiating to the circumference. This fossil may represent an equisetaceous nodal diaphragm. This fossil compares with the fossils from the Rhaeto-Liassic of Greenland which Harris (1931, p. 11-12) called “star-caps’’. Similar fossils have been reported from the Late Triassic — Early Jurassic of Queensland by Jones (1948, pl. 1, fig. 1) who called them “star-caps”. Such fossils also have been observed in the Middle Triassic Basin Creek flora at Nymboida by WBKH. Harris (1931) found that “‘star-caps” were borne terminally on equisetaceous axes and he suggested that they are formed from a modified leaf-sheath in which the teeth have been bent over the stem apex and have adhered together. Material. AMF 59419 from locality 1583 and UNEF 15530 from locality 1695. Fig. 7. Note: Specimens natural size unless stated otherwise. Those photographed under kerosene are marked with an asterisk (*). 1-2 Dicroidium voisey: sp. nov., *, holotype, AMF 59447. 2, Specimen showing lateral venation. Paratype. AMF 59448, x4. 3-5 Voltziopsis townrovi sp. nov., *, Female cone. 3, AMF 59449, holotype. 4, AMF 59450. 5, AMF 59451. 6 Voltztopsis sp., *, Foliage shoot, AMF 59452, X3. 7 Taenzopteris lentriculiforme (Etheridge) Walkom, AMF 59453. 8 Rhipidopsis ? narrabeenensis Walkom, AMF 59455. 9 Dispersed seed a, AMF 59454, x2. 10 Stem ?, AMF 59456. Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 68 AN EARLY TRIASSIC MEGAFOSSIL FLORA CONCLUSION On the basis of common forms, the Lorne Basin flora probably correlates best with that occurring in the Early Triassic Upper Newport Formation of the Sydney Basin. There are few forms which occur also in the Anisian-Ladinian floras described by Retallack (in Retallack, Gould, and Runnegar 1977) from the Clarence-Moreton Basin. On palynological evidence Helby (1973, p. 146) suggests that the Camden Haven Group is probably late Early Triassic in age, a determination which we believe is supported by the macroflora described in this paper. Pratt and Herbert (1973) reconstructed the Lorne Basin palaeoenvironment as being an area of alluvial fans debouching from the south-west, west and north-west onto a widespread plain crossed by meandering streams. An arid climate was suggested. The localities from which the bulk of our flora was collected all represent a single environment, that of back-swamps bordering streams and filled by overbank flooding. ACKNOWLEDGEMENTS We thank Dr R. E. Gould, Dr B. C. McKelvey and Dr G. J. Retallack for much helpful discussion. We very much appreciate Dr Retallack allowing us unrestricted access to his unpublished thesis on the palaeobotany of the Newport Formation. The curators at the Australian Museum, The Mining and Geological Museum, Sydney and the Universities of Sydney and New England kindly allowed access to their fossil collections. Mrs Felicity Holmes greatly assisted with the collecting, photographing and typing. Dr Gould photographed some of the specimens and Mrs Rhonda Vivian did the final typing of the manuscript. S.A. received assistance from the United States- Australia Cooperative Science Program and was provided with research facilities in the Department of Geology of the University of New England. 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The floras of the Burrum and Styx River Series. Pub. Geol. Surv. Queensl.,263: 1-75. ——, 1924. — On fossil plants from Bellevue, near Esk. Mem. Queensl. Mus., 8: 77-92. ——, 1925.— Fossil plants from the Narrabeen Stage of the Hawkesbury Series. Proc. Linn. Soc. N.S.W., 50: 214-224. ——, 1932.— Fossil plants from Mt Piddington and Clarence Siding. Proc. Linn. Soc. N.S.W., 57: 123- 126. WALTON, J., 1936.— On the factors that influence the external form of fossil plants. Roy Soc. London, Philos. Trans. B226: 219-237. WHITE, D., 1929. — Flora of the Hermit Shale, Grand Canyon, Arizona. Carnegie Inst. Wash. Pub., 405: 1-119. Proc. Linn. Soc. N.S.W., 108 (1), (1978) 1979 A natural habitat of the insect Pathogenic Fungus Culzcznomyces in the Sydney area RICHARD C. RUSSELL, MARGARET L. DEBENHAM and DAVID J. LEE RussELL, R. C., DEBENHAM, M. L., & LEE, D. J. A natural habitat of the insect pathogenic fungus Culzcinomyces in the Sydney area. Proc. Linn. Soc. N.S.W. 103 (1), (1978) 1979: 71-73. One possible source of a mosquito pathogenic fungus, Culzcznomyces sp., which appeared in a Sydney laboratory colony in 1972, was considered to be McCarr’s Creek near Sydney. After several seasons’ search, larvae of the mosquito Aedes rupestris Dobrotworsky, collected from rock pools in the creek in late 1976 and early 1977, were found to be infected with Culzconomyces. The fungus was cultured zn wtro and tested for pathogenicity. R. C. Russell, M. L. Debenham and D. J. Lee, School of Public Health and Tropical Medicine, Commonwealth Department of Health, University of Sydney, Australia, 2006; manuscript received 22 June 1977, in amended form 20 December 1977. INTRODUCTION A sudden increase in larval mortality occurring in February, 1972, in a colony of Anopheles amictus hilli Woodhill & Lee maintained at the School of Public Health and Tropical Medicine, University of Sydney, was traced to an infection with a pathogenic fungus, but attempts to identify the fungus were unsuccessful (Sweeney et al., 1973). In September, 1972, a similar phenomenon was observed by Prof. J. N. Couch and associates in colonies of Anopheles quadrimaculatus Say maintained at the University of North Carolina, Chapel Hill, N.C. In 1974 Couch, Romney & Rao described the Chapel Hill fungus as a new species, clavosporus, of a new genus Culicinomyces, stating that the fungus had been introduced to the mosquito colonies in water from University Lake, Chapel Hill, and adding that it also occurred “in mosquitoes in water from a lake near Sydney, Australia (Dr Lee and associates)”. Apart from the implication that the two fungi are conspecific — and they have as yet only been determined to be congeneric (Sweeney, 1975a) — this statement is misleading, as at that time the source of introduction of the fungus to the S.P.H. & T.M. colony was unknown, although Professor Couch had been informed that one of the suspect sources was McCarr’s Creek near Sydney. SOURCE OF THE FUNGUS A review of the situation in the mosquito colony at the time of introduction of the fungus revealed three possible sources: (1) it had persisted undetected at a low level since the colony’s development from larvae originating in Gove, N.T., twelve months previously, but had reached epizootic level following a change in pupal separation procedures ; (ii) it was introduced with Aedes australis (Erichson) which had recently been colonized in the laboratory from specimens collected in brackish rock pools at Tamarama, Sydney. (i11) it was introduced with water brought from McCarr’s Creek, French’s Forest, Sydney, which was used for culture of the dn. amictus hilli larvae. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 72 A NATURAL HABITAT OF CULICINOMYCES When temperature and salinity tolerances of the fungus were considered (Sweeney, pers. comm.), (iii) was accepted as the most likely source, and investigations of this area were begun in an attempt to find the natural habitat of the fungus. Culicinomyces is pathogenic to several species in a number of different families of Dipterous insects in their aquatic larval stage (Sweeney, 1975b); it has also been found in adult mosquitoes from an infected laboratory colony, but only rarely (Debenham & Russell, 1977). Thus, over several seasons, collections were made of aquatic Dipterous larvae, and samples were taken of water, vegetation, substrate and organic debris for laboratory examination, but until recently no evidence of the presence of Culicznomyces was detected in any of these collections. Then, on 5th January, 1977, several dead Aedes rupestris Dobrotworsky larvae, part of a collection of aquatic Dipterous larvae made at McCarr’s Creek on 23rd December, 1976, were found to be showing extensive growth of a fungus identifiable as a species of Culicionomyces. These larvae all came from a single small, shallow rock pool, with bottom organic debris over gravel, in the partially dry creek bed. This pool was only a few metres above the site from which the water for the colony is collected. The larvae had been maintained since their collection in the water in which they were collected, and kept in a laboratory separated from that in which Culzcitnomyces cultures were stored. Care had been taken to eliminate any chance of contamination from these cultures. The infected larvae were washed in 50 ml of distilled water, then plated onto nutrient agar containing 0.1% streptomycin and 0.02% neomycin. The washings were retained. Growth of Culzciznomyces and other fungi occurred on these plates, and the Culzcinomyces was selectively removed and replated. Subsequent examination showed a pure culture of Culicznomyces had been obtained, and this was transferred to nutrient agar slopes on which it is currently being maintained. On 6th January, 1977, further collections of Dipterous larvae were made from rock pools in the creek bed. Prior to this collection some rain had fallen and water had flowed over these pools, but they did not appear to have been completely flushed out. On 10th January a number of Aedes rupestrzs larvae from this collection, taken from the same pool as the larvae found infected on the previous occasion, and maintained in the same way, were also found to be infected with Culicinomyces. These larvae were washed and plated, and the fungus established on nutrient agar as before. Following heavy rains the McCarr’s Creek area was re-investigated on 25th January, 1977, but the rock pools had been completely flushed through, and very few Dipterous larvae could be collected. Examination of these, and of larvae collected here and from similar creeks in the vicinity on other occasions, has so far not produced any further evidence of Culicznomyces in this area. PATHOGENICITY OF THE FIELD ISOLATIONS Spores washed or cut from nutrient agar cultures of the field-isolated fungus were added to test trays of An. amictus hilli larvae, and the development of these compared with larvae in trays to which no spores had been added. In all cases most of the larvae in the inoculated trays were dead within seven days, and growth of Culicenomyces was observed in many of the dead larvae. Larvae in control trays developed and emerged normally. Qualitatively, the pathogenicity of the field-isolated fungus appeared comparable to that of the laboratory-derived culture. Following a suggestion by Mr M. O’Keeffe, S. P. H. & T. M., the field-isolated fungus was also established on a Nocardia medium (Tsukamura, 1969 — 10g glucose, 4 g sodium glutamate, 0.5 g KH,PO,, 0.5 g MgSO,.7H,0, 20g agar, 11 Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 R. C. RUSSELL, M. L. DEBENHAM AND D. J. LEE 73 distilled water, with pH adjusted to 7.0 by the addition of 10% w/v KOH). After establishment, fungus grown on this medium was tested for pathogenicity to mosquito larvae in the manner described above, with similar results. For additional confirmation of pathogenicity a small number of An. amictus hilli larvae were added to the 50 ml of distilled water which had been used to wash the infected larvae from McCarr’s Creek prior to plating them. After four days all larvae were dead, and one showed good growth of Culicinomyces, but the remaining larvae were too heavily contaminated with Protozoa for the cause of death to be determined. CONCLUSION It has been found that the insect pathogenic fungus Culzcznomyces exists naturally in McCarr’s Creek, French’s Forest, Sydney, and that it infects larvae of the mosquito Aedes rupestris in this area. It is not yet known whether mosgito larvae are the natural hosts of the fungus in this habitat or whether such infections are incidental to the natural cycle of the fungus. ACKNOWLEDGEMENTS We wish to thank Dr A. W. Sweeney, 1 M.R.U., R.A.A.M.C., for confirmation of the identification of the fungus; and Mr M. O’Keeffe, S.P.H. & T.M., for advice on and provision of culture media. References Coucu, J. N., Romney, S. V., and Rao, B., 1974. — A new fungus which attacks mosquitoes and related Diptera. Mycologia, 66: 374-379. DEBENHAM, M. L., and RussELL, R. C., 1977.— The insect pathogenic fungus Culicenomyces in adults of the mosquito Anopheles amictus hill. J. Aust. ent. Soc., 16: 46. SWEENEY, A. W., 1975a.— The mode of infection of the insect pathogenic fungus Culzcznomyces in larvae of the mosquito Culex fatigans. Aust. J. Zool., 23: 49-57. ——, 1975b.— The insect pathogenic fungus Culicznomyces in mosquitoes and other hosts. Aust. J. Zool., 23: 59-64. ——, Lex, D. J., PANTER, C., and Burcess, L. W., 1973. eA fungal pathogen for mosquito larvae with potential as a microbial insecticide. Search, 4: 344-345. TSUKAMURA, M., 1969. — Numerical taxonomy of the genus Nocardza. J. gen. Microbiol., 56: 265-287. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 THE LINNEAN SOCIETY OF NEW SOUTH WALES RECORD OF THE ANNUAL GENERAL MEETING, 1978 The one hundred and third Annual General Meeting of the Society was held in the Science Centre, 35 Clarence Street, Sydney, on Wednesday 29 March 1978 at 7.30 pm. The President, Professor B. D. Webby, occupied the Chair. The minutes of the one hundred and second Annual General Meeting, held 28 March 1977, were read and confirmed. REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 1977-8 Publications The Society's Proceedings were issued on the following dates : Vol. 101, Part 3 6 April 1977 Vol. 101, Part 4 28 July 1977 Vol. 102, Part 1 28 September 1977 Vol. 102, Part 2 26 October 1977 During the year a Committee of Council reviewed suggestions regarding style and format for future issues of the Proceedings. A number of recommended changes have been approved by Council. These include a new cover design, changes to the layout of abstracts and headings of papers, use of a better quality paper which will allow half tones to be placed at appropriate places in the text (and not, as hitherto, printed as plates separate from the text) and publication of the Proceedings in two double issues per volume in the hope of attracting the Book Bounty which could lead to a saving of up to 25% on printing costs. It is planned to incorporate these changes into Volume 103, the first double issue of which should appear late in 1978 or early in 1979. The LINN SOC NEWS continued to be produced quarterly to keep members informed of Society activities, forthcoming events and other items of interest. Membership During the year 5 new members were elected, 12 resigned and one died. At 1 March 1978 the society consisted of 255 Ordinary Members, 26 Life Members and 5 Corresponding Members, making a total membership of 286. Miss Kathleen English, who died in August 1977, had been a member since 1930. She was an authority on Tabanid flies and published the results of her research into their life histories and taxonomy in our Proceedings. We have recently learned of the death of Dr Hans Laurits Jensen, Macleay Bacteriologist from 1929 to 1947 and Corresponding Member of the Society since 1949. The work that he did as Macleay Bacteriologist is held in world-wide esteem. Dr Jensen contributed more than 30 papers to the Proceedings. Meetings The eleventh Sir William Macleay Memorial Lecture was delivered by Professor D. T. Anderson of the University of Sydney. The lecture entitled Natural History Today was given at the Science Centre on 16 March 1978 and was followed by a dinner. Professor Anderson’s text appears elsewhere in this issue of the Proceedings. At general meetings during the year members heard talks by distinguished speakers on a wide range of topics. On 27 April 1977, Mr David Colby of the Australian Atomic Energy Commission spoke about ‘The Uranium Debate’. ‘Discovering the Australian Desert Culture’ was the subject chosen by Professor R. A. Gould, Department of Anthropology, University of Hawaii, for his talk on 27 July. Finally, on 14 December, Dr J. A. Dulhunty of the Department of Geology & Geophysics, University of Sydney, presented ‘The Lake Eyre Story’. Two Field Days were held, the first being to examine aboriginal rock carvings at West Head under the guidance of Ku-ring-gai Chase National Park Rangers in June 1977, the second (in August) was led by Dr D. Adamson and directed to the problems of invasion of exotic plants in several natural bushland sites around Sydney. To complete the record, note is made of the very successful full-day symposium on “The Natural History of the Myall Lakes Sand Masses’. This meeting, organized by Dr P. Myerscough, was held on 12 November 1977. Library Since the appointment in July of our new Librarian, Mrs Johanne Buttigieg, the backlog of work has been brought up to date and the library is now functioning normally. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 76 ANNUAL GENERAL MEETING, 1978 Linnean Macleay Fellowship Mrs Jennifer Anderson has completed her second year as Linnean Macleay Fellow at the University of New South Wales. Her tenure has been extended for a third year. She is studying seasonal cycles of development in some Australian ladybirds (Coleoptera: Coccinellidae) . Her particular interest is reproductive diapause. Factors that induce, maintain and terminate diapause are being investigated in the laboratory, in conjunction with a seasonal field study. Species being studied include the aphidophagous Scymnodes luidigaster (Mulsant) and the mycophagous Leptothea galbula (Mulsant). Linnean Macleay Lectureship in Microbiology Dr K. Y. Cho reports that his research work in the past year has been aimed at characterizing the membrane of Azotobacter vinelandz. The membrane lipids have been analysed and the antegenecity of the membrane studied by cross immunoelectrophoresis. In June 1977 Dr Cho participated in a UNESCO training course held in Hong Kong on the cultivation of edible fungi. Staff Appointments Following the resignation of Mr A. M. Ginges from the post of Secretary to the Society in June 1977, Council appointed Mrs Barbara Stoddard to the position as from 4 July. On 25 July, 1977, Mrs Johanne Buttigieg was appointed Librarian. Office The Society now has an office near the Library on the 6th floor of the Science Centre. The Librarian is in attendance on Mondays and Wednesdays from 9 am until 1 pm and the Secretary all day on Tuesdays. The office is closed on Thursdays and Fridays. The Society's telephone number is 290 1612. Science Centre During the year the Science Centre has become recognized as a source of services to the scientific and professional community and has attracted interest as a model for others. The Earl of Shannon, chairman of the proposed Science Centre in London, inspected the building during his recent visit to Sydney and had discussions on problems related to instituting such a project. In spite of unfavourable economic conditions, the Science Centre has progressed to the stage where all the office space has been let except for a portion of the 5th floor, all shops and car-parking spaces are taken and the Secretarial Services division has doubled in size. The use of meeting rooms and facilities has increased to the point at which there are meetings held in the building on most week-nights. The restaurant on the ground floor which closed in December 1977 is expected to reopen under new management in April 1978. The income of the Science Centre now exceeds operating expenses but is not yet sufficient to meet more than a part of the interest owing on the bank loan. A notable item of decor on the exterior wall of the Auditorium on the Ist floor is a mural of appropriate design in copper created by Mr Auriel Ragus and donated by Mr E. J. Selby, a member of the Linnean Society of New South Wales who serves as one of its directors on the board of Science House Pty Ltd. REPORT OF THE HONORARY TREASURER FOR THE YEAR 1977 The Balance Sheet and the Income & Expenditure accounts of the Society’s General Account give cause for considerable concern. Again, we have suffered a deficiency, this year of $3516. Our Accumulated Funds, at over half a million dollars, might suggest that we are a wealthy Society but four-fifths of this amount is invested in the Science Centre building which has not yet reached the stage of providing any income to the Society and is unlikely to do so for some time. Our invested funds amount only to rather less than $93,000, from which we received interest of $8903. Under Expenditure, a new item Fellowship Grant appears. Council decided that the amount which it is legally entitled to pay its Fellow from the Fellowships Account had to be supplemented in this way to bring the emolument nearer the value of scholarships awarded by other organizations. The cost of printing the Proceedings is much higher this year, partly because of further increases in the printer’s charges and partly because two parts of the Proceedings for 1976 were not paid for until 1977. The new item Room Hire represents the cost of hiring meeting rooms for the Society’s functions in the Science Centre. The item Rent is for the small office on the 6th floor of Science Centre which we occupied for part of 1977 and is much less than the rent for 1976 when, for most of the year, we had rooms in Science House. Secretarial Services includes a fee of $4000 for editing and processing the Proceedings, which underlines the fact that the cost of publishing a journal is much more than the actual printing costs. The main sources of income in the General Account are the Membership Subscriptions and Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1979 ANNUAL GENERAL MEETING, 1978 77 Subscriptions to the Proceedings, Interest on investments, the surplus income from the Fellowships Account and, in 1977, Sales chiefly of back numbers of the journal. This last fortunately was higher than usual this year but cannot be depended upon for consistent results in the future. In the Fellowships Account investments yielded more Interest in 1977 than previously because of turnover of lower interest-yielding stocks to an investment earning higher interest. As we employed a Fellow for the whole twelve months no transfer was made to the Fellowships Capital Account. After paying the salary of the Linnean Macleay Fellow the surplus of $7250 was transferred to the General Account. In the Bacteriology Account the Interest received increased slightly this year. The sum of $2500 was donated to the University of Sydney towards the salary of the Linnean Macleay Lecturer in Microbiology. The deficiency of $102 simply means we donated slightly more than we received as interest during 1977 but the reserve balance is still quite adequate. The Scientific Research Fund has been augmented by Interest of $1659 and Donations of $2000; the balance now stands at $17,773. Following presentation of the Honorary Treasurer’s Report and discussion, a motion that the Audited Balance Sheets for 1977 be adopted was passed unanimously by the members present. The President, Professor B. D. Webby, then delivered his Presidential Address entitled The Ordovician Stromatoporozds. {The text of Professor Webby’s address appears in Vol. 103, part 2 of the Proceedings. | No other nominations having been received, the Chairman declared the following duly elected for the ensuing year: President: MrJ. T. Waterhouse. Members of Council: Dr D. A. Adamson, Dr A. E. Greer, Dr L. A. S. Johnson, Mr A. N. Rodd, Dr C. N. Smithers, Professor N. G. Stephenson. Auditors: W. Sinclair & Co. Professor Webby then invited Mr Waterhouse to take the Chair. Before closing the meeting Mr Waterhouse called on Dr A. Ritchie to propose a vote of thanks to the Retiring President. Dr Ritchie made special reference to Professor Webby’s efforts while President to update the format of the Proceedzngs: his vote of thanks was carried by acclamation. Proc. Linn. Soc. N.S.W., 103 (1), (1978) 1979 ANNUAL GENERAL MEETING, 1978 78 19° 182 LOS$ 66°G06 904 66 096 36 00°616 I 00°T 00° S6r 00°€3F I 19° L1G’ 16°S6 L866 OO'IIS 09° F006 00°000'00+ 00°0SL'é1 0000166 00°000 02 00000 'T 00'00L'98 66 OLF 00°68F 00° +86 16 LS 16°0L9'T $ sta ll fescre semana nay ata tone weit SJossy JUSIIND [BIO] Ey RY SRSA Sa CHER tT OE ORR Coe ee Ce yueg 3e yse CEE AMER UA TIL cee EME puezy uo yse5 “++ yunooosy sdiysmoyay Avapey UeouuTT clo) 5-000 00-6 Hau 2o OOO mh ao D200" Gnd 2d s101q9q Arpung pape anied Sek Acbie cekmcs salts ek ope ee sae junosdy JUsWIaseUR, spabaes bepoataaes (aaa GR age ea JUNODIY 21]U9D adUaIIg —paiuiry Aig asnopy auaI9g — sueoy —Sjassy qUoLIND SOL aC ou Mo Nrr en Et Heea As SatcSINTONED RES pieog aBeureig ® adeiamas ‘1918 UeN[odonap, Dace roe peiunry ‘oD Suruyoy 1e8ng [etuojoD yeh nese hue dnvaareame see race open eat AN pour, “Or a'O sop IW — soinjueqeq EON pea) Se Cae sop JW — suvOT YI[eamUOUIUIOTZ RE RIOOS ae dion iwauidojaaaq sadimosay uelpensny SIGS Bint 5.810050 Ajato0g Zurp[ing JuoueULieg “\'S'N —susodaq — SJUIUIISOAUT ion a ao eee Ome dro Mis oso siassy paxly [210 0 nt0.b.0 0 OOS Oop peiuwiry Ag asnop 2dUsI9g — s1eYS — 3s0D IY — WOo9UW}ssAUT satis sensevachons msn es jjO U9}1IM UONeID2Id aq ssaT ROG A AL OLOmG OB Ost Ovo MoE 3809 TW — saulyoeW 230 Saree asia ee ee Re jjo uadqtim uoTeIda1daq ssaT See eee sop IW — s8uIIILy 2 samIxiy ‘amnqUINng —slassy poxly 690'909$ 19° 184 LOS$ 080' LO 0L9'S 101 b0E S006 000°00% 098°S6 OGL'31 0012 000° 000°T 00°00L°9§ o1s'8 00°966 I 99° SFL I G8 °Z0h 00S 8Eg'l Il 919° 96816 60S $ $ 9L6L 290'909$ SUA Malo OeAd O18 so BO pay suoneu0g —poiunry Aig asnox a9uaIdg¢ SCE OECECn CEC ECnCnCECnC nrc mCmnatinc Surpurqyoog SPL I — SdALISIY LL61 Tequisseq Is[g “eoueleg 616 60S Sosa pean ieak ay3 10} Aduatoyap ssaT BOER LL6I ‘Arenuef sp ‘sourjeg —spuny paiejnumsoy 9L61 LLOL ‘19qQUI999q ISTE 1¥ sv J99Yg sDULTeg LNNOOOV IVYANAD SHIVM H.LNOS MAN AO ALAIDOS NVANNIT Proc. 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N.S.W., 103 (1), (1978) 1979 ANNUAL GENERAL MEETING, 1978 82 “BLEU “YIP YIZ “IoInseoily] “UOP{ ‘AUAIOIA “M ADAOL “BL61 “YDIRW Jo Aep yquanas siya Aoupdg Ie Ga.L VC ‘popuoure se ‘GhEL ‘DOV UONesay sJUeIUNODIY dI[qng IY} JepuN paiaIssay syuejUNO DY palaeYyD “OD 9 YIVIONIS “M “Aqa190g aYI JO syoog ayy Aq pasopdsip se puk sn 0} usAIS suoeUe|dxa at) 03 SUIpI0INe 1/6] ‘1equissq STE Ww se UNIV puny YOIvasey IjJMUSIIG aya Jo sireyye [BIOULUTY 24} Jo UONIsod ay) YII0} sias A[9a1I09 Jaays sduR]eg aA0qe ay IVY) UOIUIdo oy) Jo ore puke 116 19quI290q 1ST ¢ Pepus syIWOUL dATIMI OY} IOF safe YING MIN] JO AlaI90g URUUTT ay) JO sp10daI puke syoog ay} paipne savy aM LaOdada SYOLIGNV 69 SLL LIS PIL Fg 69° ELL LI$ PLI F1$ PAG MORO HSOAuOOOUogonso oben beNE cde dhondno sob acl. yueg 3 yseg 66 —Ssjassy Juoiiny 00° 16L‘L1 160'F1 00°002‘T OOPS 6.00601) 000-60 6006. 858 ‘0 6°G'66-9)0-0 1b poiuwny sourida0y jeninyy 002‘ QO 00 Cie tere area arin eviceees Ae py] uoneiodi0Z asuridasoy je1isnpuy 006 I MMUNST BP egeaboosvooebbocesoounbensusndoenouen pr] sutpary yerauas ie COUP 4 Wig (Gentes ae Coa nce aan re ich Sate peru] uonerodi0D suoiem — “¢'D'N'd 1201 {OYIVELOYOY) EN rte sila hae NG ORO Bra OOo Deco Do PT eyersny jo uonesodioy soueury OOl‘s 00°00L°% payuiry eoueidassy [eiouey 2 [eI1auIWIOD 00L'3 GORGL TD EA eee srs mee ea tee Tiny rset gece aa te SB LL61 ‘19qQUIE99q ISTE ‘souLTeg PIL FL 00°006'T a ! ee PIT eyensny yo Auedwoy umsjonag ysnug 0026'1 Sale (OURO US Sree ene ears amt ae eno any Sree a pr] uone1odi0D aoueury [eIyaueg 009°¢ OOL000; Se ae ans PeaAtsooy suoneUuog OOOO Faint ea ere ee eee a acer pi] uonerodioy sajueieny uelensny — REE TIS. Ripe PSO BPO SG Si0\0'aD'o o.057 0 O'0.0 8,915.8 Pearaday IS919}U] —1s05 1y — soinjueqeq PP PLL Fl PSD, Oo) CER ICH OL CNBR 88) O08) 6 LL6U ‘Arenuef sy] ‘aouejeg — SJU9UI}SIAUT —spung palemumsoy $ $ $ $ $ 9L61 9L6L LLGI ‘t9quIAa99q ISTE Ie sv JIQYg sDUL[eEg JUNODY puny YI1VssayY I1JWUIIIG SHIVM HLNOS MAN AO ALAIDOS NVANNIT Proc. LINN. Soc. N.S.W., 103 (1), (1978) 1975 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 103 PARI Z PRESIDENTIAL ADDRESS The Ordovician Stromatoporoids B.D. WEBBY Department of Geology & Geophysics, University of Sydney (Delivered 29 March 1978) Syno pszs Stromatoporoids are sessile, dominantly Early-Middle Palaeozoic, frame- building, calcareous fossils, probably nearest to present-day sclerosponges or hydrozoans. The Ordovician representatives, which formed important constituents of the earliest coral-stromatoporoid-algal patch reef and carbonate bank communities, include members of three markedly different families — the Labechiidae, the Clathrodictyidae and the Cliefdenellidae. The labechiids are the most diverse and abundant Ordovician group with some nineteen genera, and a maximum development in the Middle-Late Ordovician. The clathrodictyids are a predominantly Silurian family, but three genera had already appeared in the preceding Ordovician. The cliefdenellids are restricted to one, exclusively Ordovician, genus. A systematic review of all the constituent Ordovician genera is presented. The distribution of the genera is depicted in terms of the known Ordovician stratigraphical ranges and geographical spread. The labechiids exhibit a strikingly rapid period of diversification in the Middle Ordovician (in North American stratigraphical terminology, between Chazy and ‘Blackriver’ times). Most genera appear to have dispersed fairly rapidly along the band-like Ordovician ‘equatorial’ belt, but a few like Pachystylostroma and Aulacera seem to have migrated much more slowly, having taken until late in the Ordovician to achieve a circum-equatorial distribution. The clathrodictyids and cliefdenellids made their first appearances in the Australian region in the late Middle Ordovician (North American ‘Trenton’). Clathrodictyon and Ecclimadictyon also migrated very slowly, taking until the end of the Ordovician to attain a circum-equatorial dispersal. While the clathrodictyids may have evolved from a simple ‘vesicular’ Middle Ordovician labechiid, it is more difficult to derive the complex morphology of Clzefdenella from such an ancestor. Clefdenella is only recorded from Australia and Siberia. The supposed Cambrian ‘stromatoporoids’ cannot be positively confirmed as the ancestral stocks to later stromatoporoids. They have a temporally and geographically restricted occurrence in the Early Cambrian of the Altai Sayan mountain region of Siberia, have archaeocyathan associations and morphological resemblances to some members of the class Irregulares, and they are separated by a period of 110 million years from indubitable Middle Ordovician and later stromatoporoids. An earlier ancestry would help to explain the sharp morphological differences between individual families and to resolve the problem of the derivation of Cliefdenella, but the origins may have been in non-skeletonized stocks rather than in the supposed Cambrian ‘stromatoporoids’. The first Ordovician appearance of each family seems more likely to reflect the beginnings of skeletonization in the particular group rather than to depict the initial radiation of a new lineage. The review concludes with an account of the adaptive radiation of skeletonized Middle-Late Ordovician labechiid genera, and an outline of the alternatives for deriving the clathrodictyids and the cliefdenellids. INTRODUCTION The year 1978 marked the centenary of the introduction by Nicholson & Murie of the term Stromatoporoidea. Members of this problematical group of fossil organisms have calcareous skeletons of hemispherical, laminar, encrusting, cylindrical and ramose growth form, and are composed internally of a meshwork of plate-like elements (cysts or laminae) and intersecting, vertical, rod-like pillars, or less Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 84 THE ORDOVICIAN STROMATOPOROIDS commonly, an amalgamate network. Nicholson & Murie (1878) reviewed the structure and affinities of stromatoporoids — published incidentally in the journal of our kindred society, the Linnean Society of London — arguing against their affinity with marine plants (“Nullipores’), Foraminifera, hexactinellid sponges, Bryozoa and corals. They noted the close resemblances to hydrozoans and to calcareous sponges, favouring a grouping with the sponges. Later, Nicholson (1886a) revised this view in favour of interpreting stromatoporoids as hydrozoans rather than sponges, having become convinced of the existence in various typical stromatoporoids of ‘tubes’ or ‘cells’ which may have served to house living zooids as in a coelenterate colony. Both Nicholson & Murie’s earlier interpretation and Nicholson’s later grouping have their modern adherents. Stearn (1975) has summarized the divergence of opinion as between ‘those who maintain that they are hydrozoans whose closest living relative is Hydractenza and those who believe that their closest living relatives lie in the Phylum Porifera. In favouring the assignment of them to a separate subphylum of the Porifera (the Stromatoporata), Stearn (1972; 1975) noted the morphological similarities, especially the presence of astrorhizae (possibly representing the excurrent canal system as in a sponge), and the principal difference in lacking siliceous spicules. He interpreted the stromatoporoids like modern sclerosponges as secreting a skeleton of fibrous aragonite and living as individual (non-colonial) filter feeders. On the other hand, Birkhead (1967), Flugel & Fliigel-Kahler (1968), Sleumer (1969), Mori (1969; 1970), Bogoyavlenskaya (1969), Kazmierczak (1971) and Bolshakova (1973), among modern specialists of the group, prefer to view them as having an affinity with hydrozoans. Kazmierczak (1976), though, in the latest expression of his views, radically departs from his earlier interpretation, and now proposes that stromatoporoids be removed fron the animal kingdom, and included with stromatolites in the Cyanophyta (blue-green algae). Thus the precise affinities and systematic relationships of stromatoporoids are now little closer to being settled than they were in Nicholson’s day, despite the enormous increase in knowledge of the group, including the especially important contributions on microstructure (Stearn 1966; St. Jean 1967) and soft tissue reconstructions based on analogies with living sclerosponges (Stearn 1975). The majority of past and present-day specialists have viewed the Stromatoporoidea as being a reasonably homogeneous group. However, a small band of European workers, notably Heinrich (1914), Tripp (1929), Kuhn (1927; 1939) and Alloiteau (1952), have considered them to be heterogeneous, with representatives of the family Labechiidae taken to be a separate subdivision not allied to stromatoporoids proper. These workers maintained that the coenosteum (or skeleton) of members of the order Stromatoporoidea in exhibiting a meshwork of open, gallery- like passages and a ramifying, tube-like astrorhizal system, differed fundamentally from labechiids which typically have a closed vesicular form and lack astrorhizae. The labechiids were therefore assigned to a separate order. The majority of present-day specialists of the group (Yavorsky 1962; Nestor 1964; 1966a; 1966b; Stearn 1966; Fliigel & Fliigel-Kahler 1968; Webby 1969; Bogoyavlenskaya 1969; Mori 1969; 1970; Bolshakova 1973; Khalfina & Yavorsky 1973; Khromych 1974a; and Kapp & Stearn 1975), however, include the labechiids in the Stromatoporoidea. As Nestor (1966a) has said, ‘the differences between the vesicular and other stromatoporoids are by no means so sharp and fundamental as to justify assigning them to different orders’. Many previous workers have alluded to the difficulties of interpreting — more specifically identifying — stromatoporoids because of their susceptibility to alteration both by the processes of diagenesis and recrystallization. Stearn’s (1972; 1975) explanation that stromatoporoids initially secreted a more unstable aragonite skeleton Proc. Linn. Soc. N.S.W., 108 (2), (1978) 1979 B. D. WEBBY 85 recalls Nicholson’s (1886a, p. 35) original statement. He wrote: “There is, in fact, considerable reason for concluding that the skeleton was originally composed of arragonite, and that in almost all, or perhaps all, specimens which have not been silicified, the arragonite has become more or less extensively replaced by calcite’. Other calcareous fossil groups such as brachiopods, bryozoans and some corals (notably heliolitids) usually exhibit better preservation than the associated stromatoporoids (Kapp & Stearn 1975). Not only are there problems of interpreting the effects of secondary alteration of the original structure, but there are difficulties in recognizing the limits of the original variability within many fossil species. For instance, within a single coenosteum exhibiting alternate banding of two distinctive morphologies, one may be inferred to be more completely calcified than the other (see further discussion p. 87, and Yavorsky, 1961, pl. 19, figs 1-6; pl. 20, figs 1-2). The coenosteum may have been better calcified at certain periods of growth than at others (Kapp & Stearn 1975). The first, indubitable representatives of the Stromatoporoidea appeared in the Middle Ordovician (Chazy) of eastern North America (Galloway 1957; Kapp & Stearn 1975), and the group rapidly attained a wide distribution through Europe, Asia, North America and Australia. The position of the problematical, Early Cambrian stromatoporoids from the Altai Sayan fold belt of south west Siberia (Khalfina & Yavorsky 1967) remains in doubt (see later discussion p. 112). The Middle-Late Ordovician stromatoporoids accumulated in carbonates mainly in isolated occurrences, small clusters or bank-type deposits (biostromes) . In a few places they are also reported as constituting a conspicuous component of patch reefs (bioherms) , for instance, in the early Middle Ordovician Chazy Group of Vermont (Kapp 1975), in the late Middle Ordovician Mjgsa Limestone of Norway (Skjeseth 1963), and in the Late Ordovician (Richmond) of Anticosti Island (Bolton 1972; Copper, 1974). The Ordovician stromatoporoids merit close attention for a fuller understanding of the structure and development of the early members of the group. In an attempt to maintain uniformity of usage throughout this review, I have followed North American practice of regarding the Whiterock, Chazy, ‘Blackriver’ and ‘Trenton’ as Middle Ordovician, and the Eden, Maysville and Richmond as Late Ordovician (see Fig. 8). Specimens illustrated in Figs 1-7 are from the following repositories: palaeontology collection of Sydney University (SUP), geology collection of the University of Tasmania (UTGD), Paleontologisk Museum, Oslo (PMO), and the Sedgwick Museum, Cambridge (SM). SYSTEMATIC REVIEW In general Ordovician stromatoporoid faunas tend to be less well preserved, and less diverse and abundant than their Middle Palaeozoic counterparts. Representatives of only three families have been confirmed as occurring in Ordovician strata (Webby 1969; Webby & Morris 1976) — the Labechiidae Nicholson 1879 being the largest and most varied family with its maximum development in the Middle-Late Ordovician, the Clathrodictyidae Kihn 1939, a relatively inconspicuous component of the Middle-Late Ordovician fauna, and the Cliefdenellidae Webby 1969, limited to a single, exclusively Ordovician genus. (a) Family Labechudae The conception of the family adopted herein is similar to that given by Galloway (1957), Yavorsky (1962) and Stearn (1966) — it appears to be a natural grouping, and is not easily amenable to subdivision. The group could be enlarged to superfamily Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 86 THE ORDOVICIAN STROMATOPOROIDS rank but is still not easily subdivisible. It constitutes the elements of ‘Gruppe 1’ of Fligel & Flugel-Kahler (1968). The family includes a wide range of forms characterized by imperforate, compact tissue, cyst-like to laminar plates, sometimes exhibiting denticles, sometimes pillars (occasionally both) , and with very rare obscure occurrences of astrorhizae (Yabe & Sugiyama 1930; Nestor 1966b; 1976; Mori 1970). Some workers (Kuhn 1927; 1939; Lecompte 1956) have separated the cylindrical to branching forms from the rest of the group, placing them in the family Aulaceridae Ktthn 1927. However others, such as Bogoyavlenskaya (1969; 1971) and Khalfina & Yavorsky (1973), regard the shape of the coenosteum as a feature of generic not family rank. Bogoyavlenskaya (1969; 1974) has raised labechiids to the level of an order, and Khalfina & Yavorsky (1973) and Nestor (1974; 1976) to that of a superfamily. New family subdivisions have been added — Stromatoceriidae Bogoyavlenskaya 1969, Plumataliniidae Bogoyavlenskaya 1969, Rosenellidae Yavorsky zm Khalfina & Yavorsky 1973, and Platiferostromatidae Khalfina & Yavorsky 1973 — but all seem to be unnatural divisions and should be rejected. Also, Khalfina & Yavorsky assign without comment the family Stromatoceriidae (genus Stromatocertum Hall) to stromatoporaceans rather than labechiaceans. The erection of the family Lophiostromatidae Nestor 1966 (superfamily Lophiostromatacea Nestor 1974) seems to be another unnecessary subdivision of the labechiid group, introduced in order to accommodate those elements with more massive tissue (the genera Lophiostroma Nicholson 1891 and Dermatostroma Parks 1910). 5 Nestor (1974) proposed a four-fold subdivision of the superfamily Labechiacea — families Labechiidae, Stromatoceriidae, Aulaceridae and Plumataliniidae. In a subsequent paper, Nestor (1976) offered a different four-fold subdivision adding the Rosenellidae and subtracting the Stromatoceriidae. He radically changed the generic composition of the various families from his earlier subdivision. For example, the Aulaceridae which formerly included only the cylindrical forms (such as Aulacera, Stnodictyon and Cryptophragmus) was modified to include Cystzstroma Etheridge and Stromatocerzum Hall, and to exclude Cryptophragmus and Sinodictyon — to group together the cystose forms with ‘hollow’ pillars irrespective of their growth form. The ‘hollow’ pillars of Stromatocerium, as Kapp & Stearn (1975) have remarked, may form by diagenetic processes from solid ones, and therefore may have little or no taxonomic significance. Khromych (1974a; 1974b) has given as the basis for subdividing the labechiids, the presence or absence of continuous pillars with an inverted cone-like structure, but has actually provided a most arbitrary and artificial subdivision of the group. Clathrodictyids are associated with forms like Stromatocertum (Stromatoceriidae) and Cystostroma (Cystostromatidae Khromych 1974a) in the same superfamily (Cystostromacea), and representatives of the family Labechiidae (restricted) are placed with members of the family Actinostromatidae in the superfamily Labechiacea. Stromatocertum which has close ties to Labechia and Labechiella (differing chiefly in having angular to blade-like instead of round pillars) is thus grouped in a separate superfamily from Labechza and Labechiella — placed in the superfamily Cystostromacea (and associated with Clathrodictyon). This is as extreme and as unacceptable as Khalfina & Yavorsky’s (1973) grouping of Stromatocerium in the superfamily Stromatoporacea. Kazmierczak (1971) has employed a two-fold morphological grouping with subdivisions (lineages) based on inferred evolutionary trends. Morphological group Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 87 A, lineages I-IV, contain many common labechiid genera, but the individual lineage subdivisions fall well short of acceptable natural groupings. The worst example is the association of the morphologically distinctive, non-labechiid Clzefdenella Webby with the labechiid Forolinza Nestor in lineage IV. The Ordovician representatives of the family Labechiidae exhibit a wide variety of growth form. As Parks (1910) noted, they range from ‘delicate incrustations of Dermatostroma papillatum’ to giant cylindrical columns of Aulacera like those found on Anticosti Island up to 5 metres long and 350 mm in diameter (Galloway & St. Jean 1961, p. 26). They occur, however, more commonly in hemispherical to sheet-like masses — for example, the hemispherical masses of Cystzstroma donnella Etheridge from the Cliefden Caves Limestone of New South Wales attain dimensions of 280 mm across and 200 mm in height (Webby 1969), and Labechza huronensis (Billings) from the Late Ordovician of Ontario, Indiana and Ohio has a size of up to 270 mm across and 120 mm in height (Galloway & St. Jean 1961) . Some masses of Cystostroma such as C. fritzae Galloway & St. Jean from the Late Ordovician of Ontario attain larger dimensions — 600 mm across and 250 mm in height. Kapp (1974; 1975) reported the occurrence of a still larger conical to columnar coenosteum of Pseudostylodictyon lamottense (Seely), more than 1 metre in width and height from the Chazy Group of eastern North America. (i) Cystostroma Galloway & St. Jean zn Galloway 1957 — Kapp & Stearn (1975) questioned the validity of the genus Cystostroma based on the type species C. vermontense Galloway & St. Jean from the Crown Point Formation (Chazyan) on Isle La Motte. In a survey of some 400 specimens from the type area, the morphology of C. vermontense was only found at the bases of coenostea of Labechia prima Kapp & Stearn and some other species. However in the type material re-examined by Kapp & Stearn only the vesicular Cystostroma cyst-type structure is preserved. The problem of intergrowths of two distinct morphologies in one coenosteum is highlighted by another example from the Late Ordovician of the Kolyma Basin, north-eastern U.S.S.R. Identified by Yavorsky (1961) as an intergrowth of Labechza mirabilss Yavorsky and Cystostroma rarum Yavorsky this labechiid exhibits alternating phases of growth with cysts only, giving a Cystostroma morphology, and with both cysts and pillars producing a Labechza-type structure. In another example of an undescribed species from the lower part of the Gordon Limestone Subgroup near Mole Creek, Tasmania, the bands of cysts alternate with sediment layers (less commonly diagenetic silt) . Some individual coenostea (Fig. 2E-F) show little trace of pillars, while others exhibit, in varying stages of preservation, solid pillars or ‘walless rods’ (Kapp & Stearn 1975). It is therefore assigned to Labechia. Many of the designated species of Cystostroma exhibit denticles, and some even show impersistent pillars, of the type formed by superposition of denticles through more than one cyst. The problem is where to draw the line between forms with denticles only and forms with impersistent pillars. The type species C. vermontense (based on the type material) and C. cliefdenense Webby seem to lack all traces of pillars, whereas Nestor’s (1964) C. estondense is recorded as having rare, short, ‘hollow’ pillars, and C. concinnum Ivanov zn Ivanov & Myagkova from the Late Ordovician of the Urals (Bogoyavlenskaya 1973) also has occasional slender, tubule- like pillars. Nestor (1976), who includes forms with impersistent pillars in his conception of the genus, has listed some thirteen species from the Middle-Late Ordovician. Little has been written on the ontogeny of species as a basis for determining stromatoporoid ancestry and phylogeny (see Galloway 1957). The basal row of cysts or laminae of the coenosteum (and the basal rows of successive latilaminae) may provide Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 88 THE ORDOVICIAN STROMATOPOROIDS some clues to the early developmental stages of a particular line. In this context, Kapp & Stearn’s (1975) observation that the structure of C. vermontense is only preserved in the bases of coenostea of Labechza prima and other species may be significant in supporting arguments that Cystostroma is the ‘ancestral’ stock. The base of the coenosteum of Labechza cf. L. pustulosa (Safford) — see Kapp & Stearn (1975, pl. 4, fig. 3) — and the base of a latilamina of L. aldonests Webby (1977, pl. 1d) similarly exhibit a Cystostroma-type structure. (ii) Pseudostylodictyon Ozaki 1938 — The type species, P. poshanense Ozaki 1938, following the definitions of cysts and laminae given by Kapp & Stearn (1975,p. 167), exhibits both cysts and laminae, but in some parts of the coenosteum it is difficult to distinguish between them. The laminae seemingly form parallel to the growth surface of the coenosteum, and represent successive depositional floors. The cysts, both large and small, fill interspaces between successive depositional floors in two main areas, on the slopes of the mamelons and in the troughs between the mamelons. Denticles may be present or absent. The problem of whether the laminae are original, laterally continuous, plate-like structures, or formed secondarily by the destruction of inclined-vertical side walls of long-low cysts in certain rows during diagenesis or recrystallization, is unresolved. At least in P. znequale Webby (1969, pl. 119, figs 1-3) its laterally continuous concentric plates appear to represent original primary laminae (Fig. 3F). Kapp & Stearn (1975) have expressed inconsistent statements bearing on the early stages of stromatoporoid evolution. On the one hand (p. 169) they claim that Labechia eatoni (Seely) with its rows of similar-sized long-low cysts arose from Pseudostylodictyon lamottense (Seely) with its pattern of predominant laminae, and on the other (p. 167), they maintain that ‘primitive’ species have large, strongly convex cysts of variable size and shape, while more advanced forms have smaller, more uniform long-low cysts, implying a more probable derivation of the Chazy Labechza from a vesicular Cystostroma-like ancestor. If Pseudostylodictyon is indeed ancestral to Labechza, why do not the bases of coenostea of Labechza exhibit a ‘laminar’ Pseudostylodictyon structure? Plumatalinza Nestor 1960 has been distinguished from Pseudostylodictyon by having a fine subreticulate tissue in the mamelon columns (Nestor 1964). However, it remains uncertain as to whether this tissue is of primary or secondary origin. If the latter, then the original vesicular plates of the mamelon columns have broken down into a secondary subreticulate tissue; and hence the genus should be regarded as a synonym of Pseudostylodictyon. In an opposed view, Bogoyavlenskaya (1969) has argued that Plumatalinia, because of its morphology of vesicular plates resembling laminae, and reticulate column network, could well be the ancestor of ‘laminar’ stromatoporoids, for instance the clathrodictyids. She introduced the family Plumataliniidae to accommodate this genus. Fig. 1. A-B, Labechia conferta (Lonsdale 1839), X 5, SUP 285 from the Silurian of Dudley, England. A, vertical section showing solid pillars protruding upward into the sediment. B, tangential section. C-F, Stromatocertum sp. nov., X 5; from the lower part of the Gordon Limestone Subgroup (Middle Ordovician) of Tasmania. C, vertical section of UTGD 94642 from the Cashions Creek Limestone of the Florentine Valley showing peripheral part of latilaminate coenosteum. D, vertical section of UTGD 94639 exhibiting diagenetically altered, sparry calcite-filled pillars. E, tangential section ot UTGD 94640. F, vertical section of UTGD 94641 showing sparry calcite-filled (replaced) pillars protruding into the overlying sediment. D-F, from the lower part of section on the south-west side of Sassafras Creek in the Mole Creek area. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 89 Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 90 THE ORDOVICIAN STROMATOPOROIDS (iii) Labechia Milne-Edwards & Haime 1851 — The internal structure of Labechia based on the type species L. conferta (Lonsdale) from the Middle Silurian of England has been fully investigated by Nicholson & Murie (1878) and Nicholson (1879), and shown to be composed of a large number of stout, vertically aligned pillars with a series of lenticular, upwardly convex vesicles filling the interspaces (Fig. 1A-B). The pillars, Nicholson (1879; 1886a) noted, protrude onto the upper surface of the coenosteum as solid tubercles (papillae) , but may show axial canals elsewhere. Because of the irregular arrangement of the vesicles (Nicholson 1886a), the coenosteum exhibits little tendency to split concentrically as in typical stromatoporoids with ‘concentric laminae’. Lecompte (1956), Galloway (1957), Galloway & St. Jean (1961), Yavorsky (1962), Nestor (1966b) and Mori (1970) offered a similar generic conception to that given by Nicholson, emphasizing the long, stout, round pillars, vesicular cyst plates and compact microstructure. The pillars may have an upwardly- directed cone-in-cone appearance, zigzag-shaped outer edges and axial canals as seen in vertical section, and concentric banding with lighter, apparently hollow centres (intersected axial canals) as seen in tangential section (Stearn 1966; Nestor 1966b). In fact the axial canals are not strictly hollow but have an infilling of sparry calcite. Only a few Ordovician species, like Labechia huronensis from the Late Ordovician of Canada (Galloway & St. Jean 1961), bear close morphological similarity to the type species, L. conferta. Other forms, with only a patchy development of pillars, and with more consistency in the regularity of horizontal cyst rows than in the continuity of vertical pillars, have been included in the genus; see, for example, Labechza eatoni (Seely 1904), L. prima, Kapp & Stearn 1975 and L. aldonensts Webby 1977. In the group including these species (L. przma group), the coenosteum has an overall appearance of more conspicuous horizontal elements than vertical (Figs 2D; 3C-D), in contrast to those closely allied to the type species (L. conferta group) with the continuity of strong, vertical pillars as the dominating feature of their coenostea (Fig. 1A-B). The L. prima group is well developed in the Middle-Late Ordovician, from the middle Chazyan onwards, and probably represents an early stage in the evolutionary development of Labechza. Stearn (1966) noted that the cysts of virtually all labechiids appear to be without pores except in the genus Forolznza Nestor 1964. He suggested that if the supposed foramina in the cysts of this genus are leached or recrystallized pillars then the genus would be difficult to distinguish from Labechia. Kapp & Stearn (1974) have added that if the ‘canals’ of Nestor (1964) are interpreted as altered pillars, then Forolinia should be regarded as a synonym of Labechia. (iv) Labechiella Yabe & Sugiyama 1930, Labechiellata Sugiyama 1940 and Tuvaechia Bogoyavlenskaya 1971 — Yabe & Sugiyama (1930) introduced Labechiella as a subgenus of Labechza but misinterpreted the flattened cyst plates Fig. 2. A-B, Stratodictyon sp. nov., from the lower part of the Gordon Limestone Subgroup on south-west side of Sassafras Creek, Mole Creek area, Tasmania. A, vertical section of UTGD 94643 showing pillars. Note coenosteum is cut by large calcite vein. B, vertical section of UTGD 94644 with banding seemingly partially due to replacement of rows of long-low cysts. Sparry calcite fills the diagenetically altered areas — the scattered former pillars (now ‘walless rods’) and the rows of cysts (in part resembling laminae). C, association of holotype of Stratodictyon ozakii Webby 1969 on Labechiella regularis (Yabe & Sugiyama 1930); SUP 26252, x 5, from lower part of Cliefden Caves Limestone at Licking Hole Creek, central western New South Wales. D, vertical section of Labechza of the L. prima type from the lower part of the Gordon Limestone Subgroup, south-west side of Sassafras Creek, Mole Creek area, Tasmania; UTGD 94645, x 5. E-F, Labechia sp. A, X 5, latilaminate coenostea from lower part of the Gordon Limestone Subgroup in cliff section on north side of Sassafras Creek, Mole Creek area. E, vertical section of UTGD 94646 showing a few traces of former pillars — now ‘walless rods’. F, vertical section of UT GD 94647, which in contrast lacks all traces of pillars. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 92 THE ORDOVICIAN STROMATOPOROIDS (laminae) of the type species Labechza serotina Nicholson from the Middle Devonian of England as rod-like horizontal elements. The rounded pillars of L. serotzna, as seen in tangential section, characteristically unite into chain-like groups (Galloway, 1957). Following the introduction of Labechzella Sugiyama (type species Labechiella regularis Sugiyama 1939 from the Silurian of Japan) in 1939, Sugiyama appears to have realized his error of using a preoccupied name, for in the errata column of his 1940 paper he substituted the new name Labechiellata Sugiyama 1940. In his next paper Sugiyama (1941) again referred to Labechiellata. The type species of Labechiella Yabe & Sugiyama 1930 and the type species of Labechzellata Sugiyama 1940 are closely similar except for the spacing of pillars; in Labechzella serotina they are close spaced and tend to form chain-like links as seen in tangential section, while in Labechiellata regularis, the vertical rod-like pillars are widely spaced. Galloway (1957) has argued that Sugiyama (1939; 1940) incorrectly claimed credit for erecting the genus Labechiella based on L. regularts Sugiyama, and that the introduction of the name Labechiellata Sugiyama 1941 (szc) could only be explained by a memory lapse on Sugiyama’s part. However, as outlined above, Sugiyama’s actions seem to be more readily explained in terms of his recognition of the error and substitution of the new name Labechiellata. Bogoyavlenskaya (1971) has added the new genus Tuvaechia (type species Labechia regularts Yabe & Sugiyama 1930, non Labechiella regularis Sugiyama 1939). Nestor (1976) in discussing the relationships between Tuvaechia and Labechia pointed out the lack of clear-cut distinctions between the shape of the vesicles, the spacing of the pillars and the stratigraphical distribution of the two forms. The differentiation between Tuvaechia and Labechiella is even more difficult. Tuvaechia, like Labechiellata is distinguished from Labechizella only by the wider spacing of the pillars. Like Labechiellata it should be regarded as a junior synonym of Labechiella. Labechiella regularis Sugiyama 1939 thus becomes a subjective junior synonym of Labechiella regularis (Yabe & Sugiyama 1930) and must be renamed; I propose Labechiella sugzyamaz sp. nov. Whether Labechiella (Fig. 3A-B) was derived from a Labechiza by flattening of cyst plates to form laminae or from a Pseudostylodictyon by superposition of the denticles to form strong continuous pillars remains in doubt. Galloway (1957, p. 393) claimed a derivation of Labechiella from Labechza. The developmental trend proposed by Bogoyavlenskaya (1971) involving evolution first of Labechia from Tuvaechia (= Labechiella), and second of Labechiella from a Labechza, does not seem to accord with present known facts about morphological relationships and stratigraphic distribution (see Nestor 1976, p. 37). (v) Stratodictyon Webby 1969 — The conception of Stratodictyon is here Fug. 3. A-B, Labechzella regularis (Yabe & Sugiyama 1930) from the lower part of the Cliefden Caves Limestone, central western New South Wales; X 5. A, tangential section of SUP 26240 from west of Boonderoo shearing shed. B, vertical section of SUP 26236 from Licking Hole Creek. Note well developed primary laminae. C, Labechia sp. B; vertical section of PMO 97117 from Mjésa Limestone south of Bergvika, Norway, showing diagenetically produced, ‘secondary laminae; X 5. D, Labechia aldonenszs Webby 1977; vertical section of paratype SM.A97446 from Stinchar Limestone near Girvan, Scotland, exhibiting incipient, diagenetically formed, secondary laminae; X 5. E, Cystzstroma donnellu Etheridge 1895; vertical section of assumed topotype SUP 28246 from lower part of Cliefden Caves Limestone at Fossil Hill, central western New South Wales, Xx 4. Note small denticles on upper surfaces of cysts. F, Pseudostylodictyon inequale Webby 1969; vertical section of paratype SUP 29134 from Clearview Limestone Member of Ballingoole Limestone (Bowan Park Group), Malachi’s Hill, central western New South Wales; X 10. Note the presence of both cysts and primary laminae. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 98 MEK Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 94 THE ORDOVICIAN STROMATOPOROIDS restricted to include only those fine textured, latilaminate forms with horizontal elements composed of regular, close-spaced rows of long-low cysts resembling incipient laminae, vertical elements consisting of denticles and small, discontinuous pillars occupying some areas of the coenosteum but not others, and seemingly lacking astrorhizae (cf. Webby 1969). The notion of Galloway (1957, p. 364) and Kapp & Stearn (1975, p. 167) that ‘primitive species have cysts which are large, strongly convex, and variable in shape and size’ and that more advanced forms have smaller cysts with a higher length/height ratio and more uniform character may be used to imply that Stratodictyon — the type species S. ozakiz Webby (Fig. 2C), and S. columnare Webby, from the Cliefden Caves Limestone of New South Wales, an undescribed species from the Gordon Subgroup of Tasmania (Fig. 2A-B), and S. valcourensis (Kapp & Stearn) from the late Chazyan of eastern North America — isa more advanced form than the representatives of the Labechia prima group. In L. prima Kapp & Stearn, L. eatoni (Seely) and L. aldonenszs Webby, the cysts are less regular in size and shape, and they are less conspicuously aligned in regular rows appearing as incipient laminae. (vi) Rosenella Nicholson 1886a — This morphologically simple genus is typified by its exhibiting variably sized, usually large, cysts, with or without denticles (Nicholson 1886a; 1886b; Galloway 1957; Galloway & St. Jean 1961). Rosenella (Fig. 5D) may be distinguished from Cystostroma by having relatively much larger and more variably sized cysts, and from Pseudostylodzctyon by lacking clearly defined laminae. (vil) Pachystylostroma Nestor 1964 — As pointed out elsewhere (Webby 1979), the selection of the type species of Pachystylostroma, P. ungerni (Rosen) from the early Silurian of Estonia, seems to have been an unfortunate choice, for it lacks indubitable pillars (see Nestor 1962; 1964). Indeed, at the generic level, it is indistinguishable from a Pseudostylodictyon, and even bears a resemblance to the type species of that genus, Pseudostylodictyon poshanense Ozaki 1938. Pachystylostroma fragosum Nestor is the only Estonian Late Ordovician form referrred to the genus (Nestor 1964); and it also lacks pillars and would seem more correctly allied to Pseudostylodictyon or perhaps to Rosenella. It is to be hoped that pillars will eventually be found in Pachystylostroma ungerni to validate this now well-accepted genus, characterized by the presence of both cysts and laminae, as well as pillars and denticles (Nestor, 1964; Mori 1969; Kapp & Stearn 1975). Nestor’s (1976) reference to the division of thick cyst walls into a ‘basic plate’ and a ‘covering plate’ as a diagnostic feature of Pachystylostroma seems totally unacceptable. The genus (Fig. 4B-F) differs from Pseudostylodictyon in having pillars, from Labechza in having laminae, from Labechiella in exhibiting both denticles and cysts, and from Rosenella in showing both pillars and laminae. Nestor’s (1976) tentative assignment of Rosenella woyuensis Ozaki 1938 to Pachystylostroma seems unjustified on the grounds that Ozaki’s (1938) type specimens from China, and the New South Wales material of R. woywensts (see Webby, 1969), Fig. 4. Pseudostylodictyon aff. poshanense Ozaki 1938; vertical section of SUP 26235 from the lower part ot the Cliefden Caves Limestone west of Boonderco shearing shed, central western New South Wales; x 5. B- C, Pachystylostroma sp. A from the Mjosa Limestone north of Bergvika, Norway; PMO 97112; xX 5. B transverse section. C, vertical section. D-F, Pachystylostroma sp. B, X 5. D, vertical section of UTGD 94649 from the middle part of the Gordon Limestone Subgroup just below the Plzomerina siltstones, Mole Creek, Tasmania. E, vertical-oblique section of UTGD 25374 from Gordon Limestone Subgroup at Gunns Plains. F, transverse section of UT GD 94648 from same horizon and locality as Fig. 4. D. G-H, new genus allied to Stromatocerium from Mjgsa Limestone south of Bergvika, Norway; PMO 97113, x 5. G, vertical section. H, tangential section. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY SORE Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 96 THE ORDOVICIAN STROMATOPOROIDS exhibit large and small cysts (but no laminae) and denticles (but no true pillars). Occasional pillars are exhibited by Rosenella amzassensis Khalfina 1960a from the late Ordovician of Gornaya Shoriya and in material from the Moiero River of northern Siberia, both included by Nestor (1976) in his conception of woyuenszs. It would seem preferable to group the Gornaya Shoriya and Moiero material in amzassensis and, despite the lack of well defined laminae, tentatively assign it to the genus Pachystylostroma. This species is not synonymous with Rosenella woyuensts. (viii) Stromatocertum Hall 1847 — The genus resembles Labechzella but the pillars have angular, meandriform, stellate or flanged outlines, as seen in tangential section. The cyst plates are frequently but not always flattened into long-low cysts or laminae. Denticles are not present. Pillars of virtually all described species of Stromatocertum appear to be secondarily altered, possibly mainly by diagenetic leaching of the interiors of the pillars, and subsequent infilling by sparry calcite. Even the upwardly protruding tips of the pillars on the upper surfaces of the coenostea are replaced and filled with sparry calcite (Fig. 1C-F). Had the pillars been originally hollow as suggested by Nestor (1964; 1976) and Kazmierczak (1971), they would not have been preserved as upwardly protruding tubercles in the sediment. To explain the formation of sparry calcite filled ‘hollow pillars’ or ‘walless rods’ requires a process of diagenetic alteration (Kapp & Stearn, 1975) probably involving selective subaerial leaching of solid pillars and subsequent infill with sparry calcite. It follows that the irregular to angular pillars of many forms may have been derived by secondary processes from large, solid, possibly originally rounded vertical elements of Labechia or Labechiella type, suggesting that Stromatocertum at best should be viewed as a form genus. Nestor (1976) has recently recommended that the conception of the genus be restricted to representatives of the S. rugosum group, and that members of the S. michiganense group should be included in a separate genus. Pillars of the first type are ‘hollow’, wall-like, of variable cross-sectional shape, usually radiating from centres which may include associated mamelons (group A of Fig. 10). Pillars of the second type are thin, complexly bent and intermeshed vertical plates, not radiating outward from centres (group C of Fig. 10). A third group, the S. canadense group, were assigned by Nestor (1976) to the genus Cystzstroma Etheridge 1895. Nestor’s S. canadense group has at least one form exhibiting denticles and therefore strictly equatable with Cystzstroma but there are others with ‘hollow’, rounded to angular pillars and no denticles which should be included in a separate group (group B of Fig. 10). A good example is the new species (Fig. 1C-F) presently under description from the lower part of the Gordon Subgroup in Tasmania (Webby, in press b) . In their classification of stromatoporoids, Khalfina & Yavorsky (1973) introduced the genus Platzferostroma to accommodate forms with Stromatocerium- like pillars and upwardly convex cysts (like the type species of Labechia). They assigned Stromatocerium huronense Billings, described by Parks (1910) from the Late Ordovician of North America, to the new genus. However, this species was subsequently revised by Galloway & St. fean-(1961) and assigned in part to Labechia huronensis (Billings) and in part to Stromatoceritum granulosum (James). The former has round pillars and the latter does not have conspicuously arched cyst plates. The former should be retained in Labechza and the latter in Stromatocerium. The genus Platzferostroma (with its type species Stromatocerium hydridium Dong 1964) is thus best regarded as restricted to the lowest levels (Etreoungtian) of the Carboniferous. (ix) Cystestroma Etheridge 1895 — Like Stromatocerzum it typically exhibits a Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 97 coarse-textured structure of large pillars and cysts, but the pillars have a rounded to irregular or serrated outline in tangential section, and tiny denticles occur on upper surfaces of cyst plates and in places on outer surfaces of the large pillars (Fig. 3E). It is likely that the large pillars of Cystzstroma were derived from mamelon columns of a form like Pseudostylodictyon poshanense Ozaki (Fig. 4A), as outlined previously (Webby, 1969). There is little likelihood, owing to the markedly different order of magnitude of size difference between the pillars and denticles, of the pillars being derived by superposition of the denticles as in other labechiids such as Labechza and Pachystylostroma (Kapp & Stearn, 1975). Stromatocertum canadense Nicholson & Murie 1878 from the Middle Ordovician of eastern North America (and perhaps the Estonian counterparts including S. sakuense Nestor 1964) should be assigned to Cystestroma. North American specimens have denticles on the upper surface of cyst plates and ‘short, spine-like flanges’ on the pillars (Parkes, 1910; Galloway & St. Jean, 1961, p. 60). (x) New genus allied to Stromatocertum — The distinctive Stromatocerium-like morphology with long, slender, ‘composite’ vane-like pillars and denticles (Fig. 4G-H) is currently being described as a new genus based on material from the Middle Ordovician Mjgsa limestone of Norway (Webby, in press, a). Pachystylostroma and the related Stylostroma Gorsky 1938 (also including forms previously assigned to Pseudolabechia Yabe & Sugiyama 1930 but now excluded because Pseudolabechza proved to be an actinostromatid not a labechiid — see Mori, 1969; 1970) both exhibit well developed, upwardly and outwardly radiating pillars within the mamelon columns (Pachystylostroma also has them in other parts of the coenosteum). In longitudinal section the pillars may be fused to form a vertically continuous, vane-like structure — see, for example, Stylostroma gracile (Yavorsky, 1957, pl. 19, figs 1-3) — while in tangential section they exhibit a stellate pattern of outwardly radiating pillars centred on the axis of the mamelon. The ‘composite’ vane-like pillars of the new genus appears to have been derived from former mamelon columns. To what extent these structures are primary, and to what extent they are modified by secondary diagenetic alteration processes is uncertain. (xi) Lophiostroma Nicholson 1891 — Nicholson (1891), Ktihn (1927; 1939), Lecompte (1956) and Yavorsky (1962) all assigned Lophzostroma to labechiids, but Galloway (1957) preferred a grouping with the actinostromatids on the basis of the thickened skeletal tissue and laminae inflected into columns and, more recently, Khalfina & Yavorsky (1973) classified Lophzostroma with the clathrodictyids. Nestor (1966b) and Mori (1970), excluded the genus from the labechiids, assigning it instead to the family Lophiostromatidae Nestor 1966b — more recently (Nestor 1974; 1976) to the superfamily Lophiostromatacea — on the grounds of it having undifferentiated skeletal tissue completely filling the interior of the coenosteum. The series of sharply undulating ‘laminae’ (possibly representing pauses of growth rather than true laminae) defines the positions of the conical pillar-like structures, seen as papillae on the upper surface of the coenosteum. The papillae in the type species of Lophiostroma, L. schmidti (Nicholson) from the Silurian of Gotland (Mori, 1970, pls 19-20), are remarkably similar in size and appearance to those of the type species of Labechia, L. conferta (Lonsdale) from the Silurian of England and Gotland. The only named Ordovician species closely resembling the type species is Lophiostroma shantungensis Yabe & Sugiyama 1930 from the Middle Ordovician of China. Its coenosteum is almost completely filled with poorly differentiated skeletal tissue, but there are traces of pillars with upwardly-directed cone-in-cone structure, and a few large, calcite-filled cysts. This species appears to be unquestionably a labechiid, possibly derived by massive thickening of its skeletal tissue from a Labechiella, or a Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 98 THE ORDOVICIAN STROMATOPOROIDS Labechia. Lophiostroma elandiense Khalfina 1960a from the Late Ordovician of the Altai region seems to represent a rather poorly preserved Labechiella. (xii) Dermatostroma Parks 1910 — This problematical, thin, encrusting labechiid typically exhibits a papillose upper surface appearance like a Labechia or Lophiostroma (see illustration of D. scabrum in Galloway & St. jean 1961, pl. 13, fig. 1), but has a less well defined internal structure. The genus seems to be restricted to the Middle-Late Ordovician horizons of North America (Parks 1910; Galloway & St. Jean 1961). The first group has a few laminae and pillars, the second, radially crystalline prisms (possibly representing much thickened pillars) and the third exhibits a structureless mass of calcite crystals. The first group may be allied to Labechiella, the second to Lophzostroma and the third is of uncertain affinities (but has a constant encrusting association on species of Aulacera). Nestor (1974; 1976) has included Dermatostroma in the family Lophiostromatidae (and superfamily Lophiostromatacea) thus separating it from other labechiids like Labechzella. In spite of its need for revision, this genus should continue to be associated with the labechiids. (xiii) Sznodictyon Yabe & Sugiyama 1930 and Ludzctyon Ozaki 1938 — Both forms come from the Middle Ordovician of China. Sznodictyon is a fasciculate to cylindrical form with large, denticled cysts axially, and rows of smaller, long-low cysts with denticles and pillars laterally. The morphology of the axial zone resembles Rosenella, and the lateral zone, a Labechza of the L. prima-L. eatoni type. Galloway (1957) included the genus Ludictyon Ozaki in synonymy with Sznodictyon, but Ludictyon has a more-or-less broadly cylindrical coenosteum with less well defined axial and lateral zones, a pattern of alternating large and small cysts in the axial zone, rows of smaller cysts laterally, and denticles in both axial and lateral zones (Ozaki 1938) . It does not seem to be a convincing junior synonym of Sznodzctyon. Apart from its broad columnar form, Ludzctyon exhibits close morphological similarities with laminar-hemispherical Rosenella, and appears to represent an earlier stage in evolutionary development from a Rosenella-type ancestor than does Sznodictyon. (xiv) Cryptophragmus Raymond 1914, Thamnobeatricea Raymond 1931 and Cladophragmus Raymond 1931 — These slender North American Middle Ordovician forms have been grouped together as representatives of a single genus, Cryptophragmus by Galloway (1957) and Galloway & St. Jean (1961), even though there are distinctive differences between them. As Raymond (1931) noted, Cryptophragmus is unbranched, Thamnobeatricea has lateral branching and Cladophragmus exhibits bifurcating branches. Cryptophragmus has an axial column with a narrow lateral zone, and an outer sheath with the structure of Labechza which is rarely seen in continuity with the axial column. It needs to be resolved whether Cryptophragmus represents the growth of one or more than one organism. Cladophragmus, on the other hand, has no lateral zone or sheath. The relationships of these slender cylindrical genera (Fig. 5C) to laminar- Fig. 5. A, Alleynodictyon nicholsont Webby 1971 from the Gerybong Limestone Member, Daylesford Limestone (Bowan Park Group) at The Ranch, Bowan Park area, central western New South Wales; transverse section of holotype SUP 34170, X 4. B, Aulacera sp. A from the uppermost part of the Gordon Limestone Subgroup at The Den, Mole Creek area, Tasmania; transverse section of outer zone of large specimen, UTGD 94651, Xx 4. C, Cryptophragmus? sp. from the Lower Limestone Member of the Benjamin Limestone (Gordon Limestone Subgroup) in the Florentine Valley; vertical section of UTGD 94654, x 4. D, Rosenella sp. from the Gordon Limestone Subgroup at Ida Bay; vertical section of UTGD 94650, x 4, showing prominent mamelon-like upgrowth. E, Aulacera sp. B from the uppermost part of the Gordon Limestone Subgroup at The Den, Mole Creek area; vertical section of UTGD 94652, x 2. F-G, Aulacera sp. C from the upper part of the Gordon Limestone Subgroup at Gunns Plains; X 2. F, transverse section. G, vertical section. Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 99 Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 100 THE ORDOVICIAN STROMATOPOROIDS hemispherical labechiids are not clearly established. They may have evolved from mamelon-like columns extending upwards off a laminar-hemispherical base, but such bases have not yet been found. They may have been derived from a strongly mammillated representative of Cystostroma, Pseudostylodictyon or Rosenella with its mamelons exhibiting large ‘axial’ cysts. (xv) Alleynodictyon Webby 1971 — The genus (Fig. 5A) appears to be more closely related to Cryptophragmus and its allies than to Sznodictyon and Ludictyon. It differs from all other cylindrical labechiids by exhibiting blade-like pillars. Representatives of Stromatocertum and the Stromatocerium-like new genus with blade-like pillars provide a clue to the possible mode of derivation of the pillars of Alleynodictyon. In the new genus, the outwardly radiating, ‘composite’ vane-like pillar structure (Fig. 4 G-H) seems to have formed from the many upwardly and outwardly inclined, small pillars fusing in continuous vertical rows and linking axially. Each such ‘composite’ pillar seems to have been derived at the axis of a former, Pachystylostroma-like mamelon column by apparent coalescence of numerous small pillars (Fig. 4 B-F). Alleynodzctyon (if the cylindrical branches are viewed as modified mamelon columns) have a differentiated coenosteum with a lateral zone and an axial column. The small pillars which are similarly formed by fusion into vertical rows are however confined to the lateral zone. They resemble septa in rugose corals. (xvi) Aulacera Plummer 1843 ( = Beatricea Billings 1857) — The coenosteum of Aulacera is unbranched, and it has a relatively wider lateral zone than in other cylindrical labechiids. The axial column is usually sharply differentiated from the lateral zone, and exhibits a series of large, superposed cysts; the lateral zone shows rows of imbricated, smaller cysts with pillars distributed, usually sporadically, in the outer part (Fig. 5B, E-G). Specimens of large dimensions have been recorded from the Late Ordovician of Anticosti Island (Twenhofel 1928). In places along the coast of Anticosti, they occur so thickly ‘as to resemble piles of petrified logs’. They protrude like hollowed trunks ‘suggesting small cannon projecting from a wall’ (Twenhofel, 1927, p. 105) in a cliff face on Anticosti, appropriately named Battery Point. Mostly they lie in the plane of bedding, but occasionally a specimen is seen to be orientated in a vertical or upright (?growth) position. Attachment bases have not been positively confirmed. Most workers (Schuchert 1919; Twenhofel 1927; Yavorsky 1955; 1957; Galloway & St. Jean 1961) have interpreted Aulacera as having had an erect life orientation, growing up vertically off an attachment base. However, Copper (1974, p. 379) has argued that Aulacera may have rolled around on the sea floor instead of growing vertically ! In preferring a vertical mode of growth, it should be emphasized that only a relatively small part of the total preserved length of the coenosteum (observed to a maximum of 5 m) — the apical growing area of Webby (1971) — may have projected above the sea floor at any one time, and upward growth may have kept pace with the adjoining sediment accumulation. However, a period of intense erosion and bodily transport of these giant columnar ‘structures’ would have been required to reorientate them into their final resting place, in the plane of bedding. Aulacera, though apparently confined to the Richmond (Late Ordovician) of North America first appears in stratigraphically lower levels in Tasmania (possibly equatable with the Eden or Maysville), and possibly from still lower horizons in China — Aulacera petchuangensis Ozaki 1938 being recorded from the Middle Ordovician (‘Blackriver’ or “Trenton’) of China. (b) Family Clathrodictyidae Galloway (1957) in an outline of the phylogeny of Stromatoporoidea regarded Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 101 the Labechiidae with their simple morphology and earliest appearance as ancestral directly or indirectly to all other stromatoporoid families. Galloway viewed the Clathrodictyidae as having evolved from a member of the labechiids in the Late Ordovician, the overlapping or imbricated cysts of labechiids being replaced by ‘the side-by-side placing of the cysts’ of clathrodictyids. A similar derivation of clathrodictyids (Jaminar’ stromatoporoids) from labechiids (‘vesicular’ stromatoporoids) through a succession of developmental stages from Late Ordovician to Early Silurian time is inferred by Nestor (1966a). He depicted a trend from a Cystostroma-like ancestor composed of imbricate, upbranched cyst plates, but not arranged in orderly rows, to Clathrodictyon vormsiense Riabinin from the Late Ordovician (Vormsi horizon) of Estonia with regular rows of long-low cyst plates, to more typical representatives of the genus, in the very topmost beds of the Ordovician (Porkuni horizon of Estonia) — forms such as Clathrodictyon gregale Nestor and C. zonatum Nestor with irregular laminae. Bogoyavlenskaya (1969) and Kazmierczak (1971) have made similar suggestions regarding the derivation of ‘laminar’ stromatoporoids from Cystostroma. It is perhaps significant that both labechiids and clathrodictyids typically exhibit a microstructure of non-porous, speckled, compact tissue (Stearn 1966). Neither Galloway, Nestor, Bogoyavlenskaya-nor Kazmierczak explained precisely how the pillars of Clathrodictyon were formed. Typically confined to a single interlaminar space (Stearn, 1966), the pillars are not as Galloway (1957) contended, strictly the downward continuations of the edges of the cysts, otherwise the intersected cyst walls would appear as circles or polygonal outlines in tangential sections (Khalfina & Yavorsky 1967). The funnel-like (or inverted cone-shaped) pillars of some species of Clathrodictyon (Stearn, 1966; Stearn & Hubert 1966) may reflect an origin of the pillars from intersections of downwardly-inflexed cyst plates. However, usually the pillars show little or no trace of associated downwardly-inflexed cyst tissue. If Clathrodictyon evolved from the labechiids then the ‘labechiid-type’ cyst walls must have broken down or failed to calcify. The regularity in the spacing of pillars may be explained by the pillars having been derived at points of intersection of former downwardly-inflexed cysts. (i) Clathrodictyon Nicholson & Murie 1878 — Characteristically Clathrodictyon is a Silurian genus with a widespread and abundant distribution (Nestor 1964; 1966b; 1976; Birkhead 1967; Mori 1969; 1970; Bolshakova 1973). Its Late Ordovician record is more restricted but not confined to Estonia and Anticosti Island as previously indicated by Galloway (1957). Nestor (1964) described some five species from Estonia, and proposed two of them as name-bearers from his Late Ordovician stromatoporoid zones. Webby & Banks (1976) have recorded an additional four species from the Late Ordovician (probably about Late Caradoc in age) of Tasmania (Fig. 6 C-D). One Tasmanian species C. zdense is notable among Ordovician forms for exhibiting dissepiments which link the pillars and subdivide the galleries. Species allied to the Estonian forms have also been reported by Webby (1969) from the Late Ordovician of New South Wales (Fig. 6A-B), and by Bogoyavlenskaya (1973) from the western Urals. (ii) Ecclimadictyon Nestor 1964 — The genus has a similar distribution and stratigraphic range to that of Clathrodictyon. It is related to Clathrodictyon but differs in exhibiting zigzag-shaped laminae, and relatively less conspicuous pillars. Although not clearly developed in all species, the pillars are confined to interlaminar spaces and tend to be alternating, not superposed; they formed from downwardly inflected laminae seemingly also at intersections of ‘cyst plates’. A distinctive orderly Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 THE ORDOVICIAN STROMATOPOROIDS Sos AS. A a. <= ~* Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 103 meshwork is represented in tangential section by the pattern of distribution of the pillars and intersecting downwardly inflected chevron-shaped laminae. Nestor (1974) has inferred a derivation of Ecclimadictyon either from Clathrodictyon or, less certainly, directly from Cystostroma. In the Estonian successions, the first clathrodictyids to make their appearance are Clathrodictyon vormstense and C. microundulatum whereas in New South Wales Ecclimadictyon nestort (Fig. 6H) first appears stratigraphically well below the first representative of Clathrodictyon (see Webby & Morris 1976, fig. 1). It comes in, indeed it defines, the base of Fauna 2, at a stratigraphical level which is no later than Early Eastonian (= middle Caradoc, or late Middle Ordovician in terms of North American or Baltic successions) . Late Ordovician species of Ecclimadictyon include E. porkunz (Riabinin) and E. kowgiense Nestor from Estonia (Nestor 1964), E. genzculatum Bogoyavlenskaya 1973 from the Urals, E. amzassensis (Khalfina) from south-west Siberia and New South Wales (Khalfina 1960a; Webby 1969), E. nestorz Webby 1969, E. cribratum Webby & Morris 1976 from New South Wales, and E. undatum from Tasmania (Webby & Banks 1976). Both the Estonian species are from the uppermost (Porkuni horizon) part of the Ordovician, and differ from the others in exhibiting dissepiments. Two other species are especially distinctive — E. amzassenszs exhibits overall vertical continuity in alignment of zigzag-shaped rows of pillars (Fig. 6E-G), and E. cribratum has patches of horizontal tissue with ‘hexactinellid’ rod-like radial processes (Fig. 61). (111) Plexodictyon Nestor 1966b — Nestor (1966b), Stearn (1969) and Bogoyavlenskaya (1973) have regarded the genus Plexodictyon as restricted to Late Silurian strata. However, P. 2?cascum Webby & Morris 1976 from the Late Ordovician of New South Wales (Fig. 7D), is a species with close morphological similarities to the late Silurian representatives of the genus, and suggests a much earlier derivation of Plexodictyon from Ecclimadictyon than previously thought. It may be significant that a Plexodictyon-like structure develops in mamelon-like upgrowths of the coenostea of some of the earliest known species of Ecclimadictyon and Clathrodictyon (see Webby & Banks, 1976, p. 131, pl. 2, fig. 5 and Webby & Morris 1976 p. 132, Fig. 5D). (c) Family Cliefdenellidae Webby (1969) and Webby & Morris (1976) have discussed relationships between Cliefdenellidae and other stromatoporoid groups. The family contains only the one genus, Cliefdenella Webby 1969. No other Ordovician stromatoporoid exhibits such a complex association of primary laminae with denticles on upper surfaces, dissepiments and a ramifying meshwork of horizontal astrorhizal canals filling interlaminar spaces, Fig. 6. Clathrodictyon cf. microundulatum Nestor 1964 from the Clearview Limestone Member of the Ballingoole Limestone (Bowan Park Group), near Malachi’s Hill, central western New South Wales; vertical section of SUP 29133, x 5. B, Clathrodictyon aff. mammillatum (Schmidt 1858) from the Davy’s Plains Limestone Member of the Daylesford Limestone (Bowan Park Group) at Quondong, central western New South Wales; vertical section of SUP 43173, X 10. C-D, Clathrodictyon plicatum Webby & Banks 1976 from the uppermost part of the Gordon Limestone Subgroup in Tasmania, X 10. C, tangential section of holotype UTGD 94626 from roadside locality leading to main quarry west of The Den, Mole Creek. D, vertical section of paratype UTGD 94629 from The Den. E-G, Ecclimadictyon amzassenszs (Khalfina 1960a) from the upper part of the Cliefden Caves Limestone at The Island, Cliefden Caves. central western New South Wales,-x 5. E-F, tangential and vertical sections of SUP 26206. G, vertical section of SUP 26207. H, Ecclimadictyon nestort Webby 1969 from the upper part of the Cliefden Caves Limestone at The Island; vertical section of paratype SUP 26200, x 10. I, Ecclimadictyon cribratum Webby & Morris 1976 from limestone breccia at the top of the Malongulli Formation, near Malongulli Trig., central New South Wales; vertical section of holotype SUP 78266, X 10. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 104 THE ORDOVICIAN STROMATOPOROIDS and vertical astrorhizal columns with associated updomed laminae, astrorhizal canals and vertical spine-like elements (Fig. 7A-C). It resembles a Plexodictyon-type structure (Webby & Morris 1976) with the addition of the complex astrorhizal system and ‘tube-like’ pillars. The vertical elements of the coenosteum — the ‘tube-like’ pillars and the astrorhizal columns — seem to retain their independence as discrete units and never appear to exhibit interconnection one with the other. Origins of this group are obscure. Cliefdenella makes its first appearance in beds of Fauna 2 (about middle Eastonian or, in North America terms, “Trenton’) of the N.S.W. succession, only about 100 m stratigraphically above the first incoming of clathrodictyids (Webby & Morris, 1976). The genus is known through a stratigraphical range in N.S.W. from the middle Eastonian to early Bolindian (Faunas 2-3 of Webby 1969), and from the Late Ordovician of Salair in south-west Siberia (Khalfina & Yavorsky 1974). STRATIGRAPHIC DISTRIBUTION (a) Australia (3) New South Wales.— Three biostratigraphically distinct coral and stromatoporoid faunas (Faunas 1, 2 and 3) have been recognized from the Late Ordovician (Gisbornian-Bolindian) limestone successions of central western New South Wales (Webby 1969; 1971; 1975). The labechiids, Pseudostylodictyon aff. poshanense Ozaki, Stratodictyon ozakiz Webby, S. columnare Webby, Rosenella woyuensts Ozaki, Labechiella regularis (Yabe & Sugiyama), Cystestroma donnelli Etheridge and Alleynodictyon nicholsont Webby appear in Fauna 1 (Fig. 8), of probable late Gisbornian age. In terms of the North American sequence, this level would be roughly equivalent to a late Porterfield? or early Wilderness age (i.e., in the older nomenclature, a ‘Blackriver’ age). This is the oldest stromatoporoid fauna known from New South Wales; and it already exhibits considerable diversity from the basic labechiid stock. The first appearance of clathrodictyids defines the base of Fauna 2, in sequence some 94 m above the top of the lower, thinly bedded member of the Cliefden Caves Limestone (Webby & Morris, 1976). On the basis of present correlations, these clathrodictyids (Ecclimadictyon nestort Webby and Plexodictyon ? sp.) probably first appeared about the beginning of the Eastonian and the first cliefdenellids in the middle Eastonian. The stratigraphic distribution of the Ordovician clathrodictyid and cliefdenellid stromatoporoids was shown in fig. 1 of Webby & Morris (1976). The labechiid component of Fauna 2 includes Cystostroma cliefdenense Webby, Pseudostylodictyon inequale Webby and Labechiella variabilis (Yabe & Sugiyama). Cystostroma cliefdenense seems to be in no way closely linked or related to the first clathrodictyids. Indeed the clathrodictyids actually appear stratigraphically below the first occurrences of C. cliefdenense. In terms of North American successions, Fauna 2 would equate with the ‘Trenton’ (Fig. 8), or in the revised terminology of Sweet & Bergstrom (1976), the Rocklandian-Shermanian. Fauna 3 spans an interval from the late Eastonian to the early Bolindian, broadly equivalent to the Eden-Maysville interval of North America (Fig. 8). The Fig. 7. A-C, Cliefdenella etheridgec Webby 1969 from the upper part of the Cliefden Caves Limestone, The island, Cliefden Caves area, central western New South Wales: X 5. A, vertical section of holotype SUP 24157. B, tangential section of paratype SUP 24154. C, oblique-vertical section of paratype SUP 24156. D, Plexodzctyon? cascum Webby & Morris 1976 from the Clearview Limestone Member of the Ballingoole Limestone (Bowan Park Group) of central western New South Wales; vertical section of paratype SUP 77277, X 7.7. Note the association with the rugosan Palaeophyllum patulum McLean & Webby 1976. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 105 ‘ Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 \ 106 THE ORDOVICIAN STROMATOPOROIDS stromatoporoids occur mainly in the upper part of the respective limestone successions at Bowan Park and elsewhere, and may represent a mainly ‘early Bolindian’ fauna. They include cliefdenellids, clathrodictyids (including Plexodzctyon? cascum) and the labechiid, Pseudostylodictyon inequale. No stromatoporoids have been found in higher Ordovician horizons of New South Wales. (ii) Tasmaniza.— Carbonates of the Gordon Subgroup accumulated through much of Ordovician time on the Tasmanian Shelf (Banks, 1962; Webby 1976; 1978). Only the clathrodictyid stromatoporoids from the upper part and labechiids from the lowest part of the sequence have so far been studied in detail (Webby & Banks, 1976; Webby, in press b). The bulk of the fauna consists of labechiids (no cliefdenellids are known), and comprises a very diverse and abundant component of the total biota. Corbett & Banks (1973) have outlined the general faunal and floral succession in the Gordon Subgroup of the Florentine Valley area as consisting of some eight stratigraphically distinct ‘faunas’, probably ranging from about Castlemainian (late Arenig) to early Bolindian. The first three ‘faunas’ lack stromatoporoids but the fourth contains Stromatocertum in an assemblage with Maclurites and Girvanella. It occurs in the Cashions Creek Limestone of probable Darriwilian age — approximately equivalent to middle-late Chazy of North America. The stromatoporoid faunas from this horizon to the top of the subgroup (from the fourth to eighth ‘faunas’ inclusive) are more complete and better preserved in the Mole Creek section than in the Florentine Valley. The first stromatoporoids to appear in the basal part of the Gordon Subgroup of the Mole Creek area are Stromatocerium, Stratodictyon and Labechia (Figs 1D-F; 2A-B, D). Stratigraphically higher, in the ‘Lzchenarza’ beds, exposed in the cliffs above Sassafras Creek, Labechza (Fig. 2E-F) is exposed in latilaminate, dome to sheet-like ‘colonies’ up to 550 mm across and 220 mm in height. Rosenella and a branching cylindrical form, possibly Cryptophragmus, make their appearance towards the top of the cliffs. Still higher the first Pachystylostroma comes in. These occurrences together with the first appearances of Tetradzum and Eofletcherza suggest a correlation with the earliest N.S.W. coral/stromatoporoid fauna (Fauna 1). Stratigraphically much higher, immediately beneath the Plzomerzna siltstones, the stromatoporoids include abundant Pachystylostroma (Fig. 4D, F) and less common Labechia, Labechiella and Rosenella. Palaeophyllum and heliolitids have been observed from this part of the sequence indicating a correlation with N.S.W. Fauna 2. Above the Plzomerzna siltstones, the faunas are rather sparse and the record of stromatoporoids incomplete. However, towards the top of the sequence of the Gordon Subgroup, at The Den and in the vicinity of the large working quarry farther west, there is a rich stromatoporoid fauna comprising Labechza, Labechiella, Rosenella, Aulacera (Fig. 5B, E) and the clathrodictyids Ecclimadictyon undatum Webby & Banks, Clathrodictyon plicatum Webby & Banks, C. molense Webby & Banks and C. sp. The fauna is associated with abundant corals including favositids, halysitids, Favest2na and other characteristic elements of Fauna 3. It is very tentatively correlated with the Eden-Maysville of North America (Fig. 8), despite the occurrence of Aulacera, which tends to be restricted to the Richmond of North America (Galloway & St. Jean 1961). Appearances of Ordovician stromatoporoid genera in Australian successions are depicted in Fig. 8. The maximum period of diversification of stocks seems to have occurred in the Gisbornian, with the appearance of Fauna 1. The species of Stromatocerium, Stratodictyon and Labechza in the lowest part of the limestone at Mole Creek are distinctly different forms (Figs 1D-F; 2A-B, D) and cannot easily be linked through intermediate forms back to a common ancestor. The species of Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B.D. WEBBY 107 Stratodictyon and Labechza may have evolved from an early Chazy representative of Pseudostylodictyon, like P. lamottense, thought by Kapp & Stearn (1975) to be the oldest North American stromatoporoid, but Pseudostylodictyon has not been found in the lowest stromatoporoid-bearing beds of the Gordon Subgroup of Tasmania. The species of Stromatocertum, has no known progenitor unless it is interpreted as having descended from Khalfina & Yavorsky’s (1974) supposed Early Cambrian Stromatocertum of the Kuznetsky Alatau region, Siberia (see later discussion, p. 113). From the initial Stromatocertum, Labechia and Stratodictyon stocks in these lowest horizons of the Gordon Subgroup, it would not have been difficult to derive the more diverse Fauna | assemblage. (b) North America Kapp & Stearn (1975) have traced the evolutionary history of the oldest stromatoporoids in North America. They have interpreted Pseudostylodictyon lamottense which first appears in the Day Point Formation (zone 1) of the Chazy Group, and has ‘denticled laminae and large irregular cysts’, as representing the earliest stage of stromatoporoid evolution. Diversification from this ancestral stock took place along two main paths, typified by the genera Labechia and Pachystylostroma. Both genera make their appearance in the Crown Point Formation (zone 2) of the Chazy Group — Labechia eatonz, L. prima and Pachystylostroma goodsellense, the first to exhibit vertical pillars supposedly secreted to strengthen the coenosteum. In the next stage of evolutionary development through the middle-late Crown Point and Valcour Formations (mid-late Chazy) a further two species of Labechia and three species of Pachystylostroma appeared, presumably derived from the main stock of L. eatonz and P. goodsellense. The genus Cystostroma which is formed solely of cysts without laminae or pillars has as its type species C. vermontense originally described from the Crown Point Formation, and considered by many workers, most notably by Galloway (1957) and Nestor (1964; 1966a), as the most primitive stromatoporoid. Unfortunately, Kapp & Stearn (1975) were unable to confirm the presence of this species and have suggested it to be a very rare species or ‘an abnormal representative of Labechza prima’. A stratigraphic break in the record of deposits and hence a gap in the course of evolutionary development occurs between the top of the Chazy Group and the base of the overlying Black River Group. According to Kapp & Stearn (1975) the basal formation of the Black River Group is the Pamelia Formation and it contains the problematical cylindrical stromatoporoid Cryptophragmus. The succeeding Lowville and Leray Formations have a new stromatoporoid fauna apparently dominated by Stromatocertum rugosum. In addition to Cryptophragmus and Stromatocerium, Galloway & St. Jean (1961) have noted the first appearance of Rosenella and the enigmatic encrusting stromatoporoid Dermatostroma Parks 1910 in horizons of ‘Blackriver’ age. With the exception of Cryptophragmus most of the labechiid genera seem to range at least to the top of the Ordovician. Cryptophragmus however is confined to the North American late Middle Ordovician (i.e., ‘Blackriver’ and possibly “Trenton’) . The large, unbranched, cylindrical stromatoporoid Aulacera Plummer is characteristic of Richmond (Late Ordovician) horizons of North America (Galloway & St. Jean 1961). Also Clathrodictyon has been reported by Twenhofel (1927, p. 107) from a similar level in the succession of Anticosti Island. The species recently figured by Copper (1974, p. 378) from a bioherm of the Ellis Bay Formation (late Richmond) on Anticosti seems to be the same species of Clathrodictyon as Dr T. E. Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 108 THE ORDOVICIAN STROMATOPOROIDS Bolton of the Canadian Geological Survey (pers. comm.) has shown me from Member 6 of this formation. It is a representative of the Clathrodictyon boreale group of Nestor (1964). Workum, Bolton & Barnes (1976) have also recorded Labechia, Cystostroma? and Aulacera from the Ordovician succession of Akpatok Island, north- east Canada. (c) Europe The oldest stromatoporoid in European successions appears to be Labechia aldonensts Webby 1977 from the Stinchar Limestone of the Girvan district of Scotland. It occurs in an horizon of approximately middle-late Llandeilo age (Williams, 1962; Williams et al., 1972). On the island of Helggya in the Nes-Hamar district of Norway, St@rmer (1953) recorded the presence of a spectacular ‘stromatoporoid reef’ some 9-10 m thick in the Mjoésa Limestone. The Mjdsa Limestone is correlated with the Upper Chasmops Limestone (stage 4bd) of Oslo, or in terms of Estonian subdivisions, equivalent to the Oandu (Dy) or Rakvere (E) stages (middle-late Caradoc). As noted by Skjeseth (1963) the core of the reef near the shore of Lake Mjdgsa consists of a massive framework of stromatoporoids. Sampling of the Mjgsa Limestone in the vicinity of Bergvika on the island of Helggya has produced a varied stromatoporoid fauna with a few individual coenostea attaining dimensions of up to 1.3 m across and 0.4 m in height. The fauna includes representatives of Labechia, Pachystylostroma and a new genus allied to Stromatocertum (Webby, in press a). To the south the Encrinite Limestone of similar age to the Mjdsa Limestone also contains stromatoporoids. Kaljo, Klaamann & Nestor (1963) have recorded two stromatoporoids from the Ashgill of Norway — Clathrodictyon microundulatum Nestor from stage 5a, and Pachystylostroma sp. nov. ex gr. fragosa Nestor from stage 5b. These elements correspond quite closely to faunas of the Pirgu (Fj.) and Porkuni (Fj;) stages of Estonia. : The Estonian Ordovician stromatoporoid succession has been fully described and analysed by Nestor (1964; 1966b). As outlined by him (1966b) the vesicular labechiids (Stromatocerium, Cystostroma and Plumatalinza) are typical of the period, but begin to be replaced by ‘vesicular-laminar’ clathrodictyids (Clathrodictyon and Ecclimadictyon) by Ashgill times. Two species of Stromatocerium are the first to appear in the Oandu (Dy) stage of middle-late Caradoc age. No stromatoporoids have been recorded from the succeeding Rakvere (E) and Nabala (F{,) stages, but the first clathrodictyids, C. mzcroundulatum and C. vormsiense appear in the next stage, the Vormsi (Fyjp,), about early-middle Ashgill time. The succeeding Pirgu (Fj,) stage contains Stromatocerium, Cystostroma and Plumatalinia in addition to C. mzcroundulatun. The topmost stage of the Ordovician in Estonia, the Porkuni (Fyy), is characterized by new species of Clathrodictyon, C. gregale, C. mammillatum and C. zonatum, by the first Ecclamadictyon (E. koigzense and E. porkunz) and by Pachystylostroma fragosum. On the western slope of the Urals, Bogoyavlenskaya (1973) has described an Ordovician stromatoporoid fauna comprising Cystostroma concinnum (Ivanov), Stromatocerium definitum (Ivanov), Clathrodictyon microundulatum Nestor and Ecclimadictyon geniculatum Bogoyavlenskaya. The species come from two horizons — the Typyl (topmost Middle Ordovician) and Rassokha (Late Ordovician, about the level of the zone of Pleurogr. linearis) . The species of Ecclimadictyon is unusual in occurring in the older Middle Ordovician (Typyl) horizon (approximately equivalent to the level of Dicranogr. clingani, or to the Oandu stage, Dy, of Estonia; Whittington & Williams, 1964) . Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 109 (d) Asza In the Moiero River section in the northern part of the Siberian Platform, Nestor (1976) has recorded a succession of Middle-Late Ordovician stromatoporoid faunas. All are labechiids, and the oldest is Cystostroma insuetum Nestor from the Krivoluk stage of possible ‘Blackriver’ (Nestor 1976, p. 9) or ?>Chazy (Sokolov et al., 1960 p. 49) age. Stromatocertum cf. sakuense Nestor 1964 occurs in an higher, Middle Ordovician (?*Trenton’) horizon, and a much larger fauna appears in the Late Ordovician Dolbor stage. This latter fauna comprises Cystostroma evenkiense Nestor 1976, Stromatocertum australe Parks 1910, S. pergratum Nestor 1976, Lophzostroma sp. and Pachystylostroma? amzassensis (Khalfina 1960) — see earlier discussion p. 96. Aulacera tenuzpunctata (Yavorsky 1955) occurs in still higher beds near the top of the Ordovician. Another form, reported by Yavorsky (1955) from the Late Ordovician of the Siberian Platform (Stony Tunguska and Moiero Rivers) , is Labechiella regularis (Yabe & Sugiyama). In the folded zone of the Altai Sayan mountain region of south-west Siberia there are many isolated records of Ordovician stromatoporoids. Khalfina (1960a) has described Labechiella lophiostromozdes (Khalfina), L. elandiense (Khalfina), Pachystylostroma? amzassensis (Khalfina) and Ecclimadictyon amzassensts (Khalfina) from the upper part of the Amzass Formation (Late Ordovician) of Gornaya Shoriya and from similar levels in Gorny Altai. In approximate terms, the upper part of the Amzass Formation correlates with the zone of Pleurogr. linearis (Sokolov et al., 1960). Khalfina & Yavorsky (1974) recorded Cliefdenella permirum from the Late Ordovician of Salair, and Bogoyavlenskaya (1971) referred to the presence of Cystostroma in the Middle and Late Ordovician of Tuva, and Labechiella regulars (Yabe & Sugiyama) and Labechza huronenszs. (Billings) from Late Ordovician horizons. From the Late Ordovician Dulankara horizon in Kazakhstan, Khalfina (1958) has described (without illustration) Labechzella kasachstanica, and from the Late Ordovician of the Chatkal Range of Kirgizia, Middle Asia, Yavorsky (1961) has recorded Cystostroma sarytschelekense. Another species of Cystostroma, C. rarum Yavorsky, of uncertain validity because of its intergrowth with Labechza mirabilis Yavorsky, is described by Yavorsky (1961) from the Late Ordovician of the Kolyma Basin of the north-eastern U.S.S.R. In addition, many species of Aulacera have been described from Late Ordovician horizons from various parts of the Soviet Union by Yavorsky (1955; 1957; 1963), in particular from the Urals, from Novaya Zemlya, from the Vilyuy and Stony Tunguska Rivers of the Siberian Platform, and from eastern Siberia. A species of Cryptophragmus, C. gracilis is also recorded from the Ordovician of eastern Siberia by Yavorsky (1955). Ordovician stromatoporoids have been reported from a number of regions of Eastern Asia — Shantung, Shansi and Liaotung (Southern Manchuria) provinces of North China, and North Korea (Yabe & Sugiyama 1930; Endo 1932; Ozaki 1938; Sugiyama 1941; Yang & Dong 1962). These occurrences all fall within the geographical limits of the major Hwangho Basin, and come from horizons of the Toufangian Series (and equivalents) of ‘Middle’ Ordovician (post Llandeilo and pre- Ashgill) age. Kobayashi (1969) has noted that stromatoporoids are the third largest fossil group in the Toufangian fauna of the Hwangho Basin, more specifically they occur mostly in horizons of the Toufangkou Limestone and the Ssuyen Formation (i.e., in the middle or upper parts of the Toufangian Series) . There is no evidence of a faunal succession — merely an abundance of elements through a relatively restricted stratigraphical range, probably equivalent to the ‘Blackriver’, and possibly the Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 110 THE ORDOVICIAN STROMATOPOROIDS | [LLANVIRNLLANDEILO CARADOC ASHGILL BRITAIN M VIRU MJUHARJU BALTIC E (stages) N WUtAZ DS /SNpILIg, juosiyoinus snjngasnijasa} $119es6 suapiqzj/nu jueBul/o s/eaul| snyeue/duioo CYSTOSTROMA : ake Vv O PSEUDO- STYLODICTYON ¥* Oa O ROSENELLA LABECHIA STRATODICTYON PACHYSTYLOSTROMA | LABECHIELLA STROMATOCERIUM CYSTISTROMA CRYPTOPHRAGMUS — CLADOPHRAGMUS — THAMNOBEATRICEA AVGIHOAEVI Bb*< view © ° q ALLEYNODICTYON LUDICTYON — SINODICTYON PCIEA GEAR MO Se NA i er hires sea Us i re are here LOPHIOSTROMA —_ — — ATT DERMATOSTROMA ECCLIMADICT YON CLATHRODICT YON PLEXODICT YON CLIEFDENELLA CLIEFDEN- Vv ELLIDAE yx NORTH AMERICA YU.S.S.R.(EXCL. ESTONIA) ACHINA CIAUSTRALIA O EUROPE (BRITAIN & BALTIC INCL. ESTONIA) AVGIALIIGOYHLV19 Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 111 ‘Trenton’ as well. The varied fauna includes records of some thirteen species from Shantung, nine from Liaotung, two from Shansi and two from North Korea. The genera include Pseudostylodictyon, Labechia, Labechiella, Lophiostroma, Rosenella, Sinodictyon, Ludictyon and Aulacera. The species of Aulacera, A. peichuangensis Ozaki, possibly represents the earliest record of the genus, earlier than the ‘Eden- Maysville’ appearances of the genus in Tasmania (Fig. 8). The association of Aulacera with other cylindrical forms (Sznodzctyon and Ludictyon) may imply an evolutionary connection, with the Chinese fasciculate, branching ‘Middle’ Ordovician forms ancestral to the unbranched Aulacera (Fig. 10). (e) General significance The Ordovician labechiids exhibit a strikingly rapid period of diversification through the interval from the Llandeilo to the early Caradoc (from Chazy to ‘Blackriver’ using North American terminology, see Fig. 8). The earliest (Chazy) stocks, judging from Australian and North American first appearances, include Pseudostylodictyon (N. Amer.), Cystostroma (N. Amer. & ?Aust.), Labechza (N. Amer. & Aust.), Stromatocertum (Aust.), Stratodictyon (Aust.) and Pachystylostroma (N. Amer.). They are followed by appearance of Rosenella (Aust.), Labechzella (Aust.), Cystestroma (Aust.), Dermatostroma (N. Amer.) and the first cylindrical forms — Cryptophragmus (N. Amer.) and Alleynodictyon (Aust.) — in the early Caradoc (‘Blackriver’) and correlatives (see Fig. 8). With the records of Lophiostroma, Sinodictyon, Ludictyon and Aulacera in the “Middle Ordovician’ of China — possibly from equivalent ‘Blackriver’ (or ? “Trenton’) levels — it seems that the maximum period of generic diversification of labechiids from basic Chazy stocks occurred early in the history of the group. They maintained their presence as the major stromatoporoid group through the Middle and Late Ordovician (from ‘Blackriver’ to Richmond times) but declined in importance from the Silurian onwards. Less than half the Ordovician genera of labechiids (and none of the cylindrical forms) survived beyond the end of the Ordovician (Fig. 8). Aulacera whose cylindrical columns attained very large dimensions (5 m in height?) in beds of the latest Ordovician was perhaps the most spectacular form to become extinct. From the apparent band-like spread of occurrences (Fig. 9), it appears that the Ordovician labechiids had approximately an equatorial distribution, perhaps limited to within 20° either side of the palaeoequator. The Middle Ordovician stocks seem to have a more restricted geographical spread — confined to eastern North America, Scotland, China and S.E. Australia — than the Late Ordovician forms (Fig. 9). Despite the inadequacies of the preserved fossil record and the unevenness in the reliability of available data, it may be suggested that some of the labechiid elements like Labechia and Cystostroma seem to have achieved an ‘equatorial’ distribution during the Chazy, while others — Pseudostylodictyon, Pachystylostroma, Stromatocerium and Aulacera — seem to have migrated very slowly to adjoining regions (Fig. 9). Others again appear to represent endemic elements, for instance, the Fig. 8. Chart showing the known stratigraphic ranges of Ordovician stromatoporoid genera. The first appearance of each genus is shown by symbols for each major region. Those genera known to survive beyond the end of the Ordovician are depicted with arrows at the top of their Ordovician ranges. Correlations between the respective British, Baltic, North American and Australian stratigraphic subdivisions are only tentative. However, it should be noted that the boundaries between the Lower and Middle Ordovician, and between the Middle and Upper Ordovician, as shown in the columns by symbols L, M and U, occur at different levels in the respective Baltic, North American and Australian successions. C/S faunas — coral/stromatoporoids faunas of Webby (1969) and Webby & Morris (1976). Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 112 THE ORDOVICIAN STROMATOPOROIDS © MID-LATE ORDOVICIAN CLATHRODICTYIDS V MID-LATE ORDOVICIAN CLIEFDENELLIDS REMNANTS OF @ LATE ORDOVICIAN LABECHIIDS GONDWANALAND 4 MIDDLE ORDOVICIAN LABECHIIDS (EXCL. ANTARCTICA) (J SUPPOSED EARLY CAMBRIAN STROMATOPOROIDS Fig. 9. Map illustrating the world-wide spread of Ordovician stromatoporoid faunas. Note also the single occurrence of the supposed Cambrian ‘stromatoporoids’ from the Altai Sayan mountain region of Siberia. Ordovician equatorial positions based on suggestions by Webby (1978) for Australia and Ross (1976) for North America. problematical, encrusting Dermatostroma of North America, cylindrical Sznodzctyon and Ludictyon of China, and cylindrical Alleynodictyon of Australia. The clathrodictyid genera Clathrodictyon and Ecclimadictyon also exhibit patterns of slow global migration like a number of the labechiid genera. The genus Clathrodictyon which first appears in “Trenton’ correlatives in New South Wales, Australia, took until the Maysville (early Ashgill) to reach Estonia, and until the late Richmond to make its first appearance in North America, on Anticosti Island. ORIGINS AND INTERRELATIONSHIPS (a) Supposed Cambrian ‘Stromatoporozds’ Although most workers have claimed the Middle Ordovician ‘vesicular’ labechiids to be the earliest representatives of the Stromatoporoidea with Cystostroma (Galloway, 1957; Galloway & St. Jean, 1961; Nestor, 1964; 1966a; Bogoyavlenskaya, 1969) or Pseudostylodictyon (Kapp & Stearn, 1975) seen to be ancestral to all later forms, a small group of Soviet specialists, notably Yavorsky (1932; 1940; 1947), Khalfina (1960a; 1960b; 1971), Vlasov (1961) and Khalfina & Yavorsky (1967; 1974), have maintained that the Ordovician stromatoporoids were derived from earlier, Cambrian stocks. The supposed stromatoporoid fauna occurs exclusively in the Early Cambrian of the Altai Sayan mountain region of south-west Siberia (Fig. 9). The fauna includes representatives of exclusively Cambrian genera (Altazcyathus Vologdin 1932; Korownella Khalfina 1960b; Praeactinostroma Khalfina 1960b; Cambrostroma Vlasov, 1961), and others which may be linked with established post- Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B.D. WEBBY 113 Cambrian forms — the genera Clathrodictyon Nicholson & Murie 1878, Rosenellina Radugin 1936 and Stromatocerzum Hall 1847. Korovinella (type species Clathrodictyon sajanicum Yavorsky) is the best known genus, typically exhibiting porous laminae, short, rod-like pillars, and vertical canals, the latter possibly analogous to the astrorhizal columns of stromatoporoids (Khalfina 1960a; 1960b). The porous nature of the laminae, on the other hand, is noted by Nestor (1966a) to ally the genus to archaeocyathans. Praeactinostroma (type species Actinostroma vologdini Yavorsky) has both discontinuous rod-like horizontal elements and pillars forming an incomplete, reticulate network also intersected by vertical canals. It appears to have closer affinities to stromatoporoids by resembling the Silurian genus Plectostroma Nestor 1964 and Mesozoic genus Actinostromaria Chalmas zn Dehorne 1920 (see Nestor 1966a). Of the other forms, Nestor (1966a) has viewed Cambrostroma as a junior synonym of Korovinella, he has noted that Radugin’s Cambrian species of Rosenellina has never been described and cannot be discussed further, and he has observed that Vlasov’s species of Clathrodictyon (C. formozovae) is based on such small fragments that they cannot be satisfactorily revised. Altaicyathus (type species A. notabilis) was originally described as an archaeocyathan by Vologdin (1932), but it was later included by Yavorsky (1940), Zhuravleva (1955; 1960), Khalfina (1960b) and Vologdin (1966) in stromatoporoids. Korovinella Khalfina 1960b has been regarded by Vlasov (1967) as a junior synonymn of Altazcyathus Vologdin 1932 (see also Hill, 1972, p. E142, and Fligel & Filiigel-Kahler, 1968, p. 525), so perhaps both Korownella and Cambrostroma should be included in Altazcyathus. However at least one species of Korownella, K. edelsteznt (Yavorsky), has recently been equated with the archaeocyathan Abakanicyathus karokolensts Konjuschkov — see Zhuravleva & Miagkova (1974, pl. 2, fig. 2). Two species of Stromatocertum, S. pospelovt Khalfina zn Khalfina & Yavorsky 1974 and S. cambricum Khalfina zn Khalfina & Yavorsky 1974 are recorded as coming from the Cambrian of the Kuznetsky Alatau, south-west Siberia. With their meandriform pillar structures, the species seem to have more in common with forms from the Early Carboniferous of China — like Stromatocerium kwangsiense Dong 1964 — than with typical Ordovician species of the genus (Parks, 1910; Galloway & St. Jean, 1961). Galloway (1957) noted that the supposed Cambrian stromatoporoids of Yavorsky (1932) were not composed of simple cysts as would be expected if they were ancestral to Cystostroma and Pseudostylodictyon but included more complex structures with laminae, pillars and astrorhizae. He implied (see also Galloway & St. Jean, 1961, p. 7) that the forms were collected from younger horizons than the Cambrian. Galloway also thought that the stromatoporoids as a group may have evolved from the archaeocyathans, from a form like Exocyathus Bedford & Bedford, but did not, in view of his doubts about the stratigraphic position of Yavorsky’s finds, explore the possibility of the supposed Cambrian forms being transitional between archaeocyathans and stromatoporoids. The more recent work of Yavorsky, Khalfina and Vlasov has proved beyond doubt that the bulk of the forms come from the Early Cambrian of the Altai-Sayan mountain region; typically they have an association with archaeocyathans, even within the one specimen. Nestor (1966a) interpreted the supposed Cambrian stromatoporoids (excluding the recently reported species of Stromatoceritum) as archaeocyathans being only convergently similar to stromatoporoids. According to him, the forms disappeared from the stratigraphic record towards the end of the Early Cambrian and had no Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 114 THE ORDOVICIAN STROMATOPOROIDS successors in the Middle-Late Cambrian or the Early Ordovician. He argued against them being the ancestors of the Middle Ordovician stromatoporoids, the earliest being the ‘vesicular’ labechiids. On the other hand, archaeocyathan specialists such as Vologdin (1966) and Hill (1972) have excluded the supposed Cambrian ‘stromatoporoids’ from a grouping with archaeocyathans. Khalfina & Yavorsky (1967) in reply to Nestor reiterated that the Cambrian forms were the oldest stromatoporoids, and that they were morphologically distinct from archaeocyathans. They noted that vertical canals were shown to be arranged like the astrorhizal canals of stromatoporoids, and the external and internal walls of typical archaeocyathans were lacking. The coniform-cylindrical ‘coenostea’ of forms like Korovinella and Praeactinostroma were seen to bear similarities to cylindrical coenostea of some later Palaeozoic (post Ordovician) stromatoporoids, for example, in exhibiting similar patterns of laminae arranged transversely to the axial canal (Paramphipora) , lateral processes (Idzostroma, Stachyodes) and a single axial canal (Amphipora, Idiostroma). The genus Clathrodictyon was seen by Khalfina & Yavorsky to be unrelated to ‘vesicular’ stromatoporoids, and possibly to have been derived from Cambrian ‘stromatoporoids’ with perforate laminae. The porous nature of the laminae in Korovinella was regarded as being different from the arrangement of pores in tabulae of irregular archaeocyathans (Suborder Archaeosyconina), and the mode of lamina formation in Korovinella similar to that of stromatoporoids like Actinostroma (another post Ordovician form) . I question these latter assertions. First, the perforate laminae of Korovinella closely resemble the porous tabulae of irregular archaeocyathans like Hupecyathus Debrenne 1964. Secondly, the perforate laminae of Korovinella differ markedly from the laminae of Actinostroma (composed of an hexactinellid network of radial processes — see Stearn, 1966, p. 86). Zhuravleva (1970) has noted that the skeletal structures of archaeocyathans of the class Irreguiares with ‘colonial’ form may be practically indistinguishable from stromatoporoids like Clathrodictyon (Zhuravleva & Miagkova, 1974). The tabulae, vertical rods and central cavities of irregular archaeocyathans are recognized as having equivalents in the laminae, pillars and astrorhizae of stromatoporoids. Zhuravleva & Miagkova (1974) have emphasized the morphological similarities between stromatoporoid Amphipora and irregular archaeocyathan Protopharetra, and between Korovinella and Archaeosycon. They have additionally shown the supposed Cambrian ‘stromatoporoids’ Korovinella edelsteint and Praeactinostroma to be indistinguishable from irregular archaeocyathans Abakanicyathus karakolensis Konjuschkov and Claruscyathus cumfundus (Vologdin) , respectively (Zhuravleva & Miagkova, 1974, pl. 2). In terms of presently accepted views of morphology and classification of stromatoporoids the Cambrian ‘stromatoporoid’ genera of Yavorsky, Khalfina and Vlasov include three (possibly four) stocks, each of which may be grouped in a different family. Altazcyathus (= Korovinella) belongs to the exclusively Cambrian family Korovinellidae Khalfina 1960b, Praeactinostroma should probably be included in the family Actinostromatidae Nicholson 1886 (a group which has no confirmed Ordovician record) and Clathrodictyon is a member of the Clathrodictyidae. A possible fourth is the presumed Stromatocertum (family Labechiidae). The korovinellids are the best known group but have the least close resemblances to Middle Ordovician or later stromatoporoids. Praeactinostroma bears no close relationships to Middle-Late Ordovician stromatoporoids. Clathrodictyon and Stromatocerrum remain too inadequately documented. Clathrodictyon is based on totally insufficient material and Stromatocertum has yet to be confirmed as coming from undoubted Cambrian horizons. Their essentially localized occurrence with archaeocyathans in the Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 115 Altai Sayan mountain region of Siberia and morphological resemblances with members of the class Irregulares, and their physical separation by such an enormous break in continuity of record of some 110 m.y. from Middle Ordovician stromatoporoids, make it impossible to establish them as the earliest, indubitable stromatoporoids. The occurrences of Clathrodictyon and Stromatocertum may represent the earliest records of clathrodictyids and labechiids, respectively, but they may equally well be viewed as homeomorphs of later stromatoporoids (either offshoots of irregular archaeocyathans, or members of an independent group). There is no record of such structures after the extinction of the reef-forming archaeocyathans at the end of the Early Cambrian, until the Middle Ordovician when skeletal reef habitats reappeared. (b) Outline of Evolutionary Development of Ordovician Stromatoporozds The Middle Ordovician stromatoporoids formed an important ‘skeletonized’ constituent of the earliest ‘coral-stromatoporoid-algal’ reef communities (Pitcher, 1971; Heckel, 1974; Copper, 1974; Kapp, 1975). Their skeletal remains were among the more prominent components of Middle-Late Ordovician patch reef and carbonate bank deposits. The advent of skeletonization may have represented an adaptive breakthrough allowing for a dramatic increase in size (some Chazy stromatoporoids are up to | m in width and height), providing the necessary support for the mantling soft tissues, and elevating the individuals above the substrate to facilitate their filter feeding (Stearn, 1972), epifaunal mode of life. The sessile stromatoporoid animal evidently lived in a moderately competitive, near-equatorial, shallow-water patch reef or carbonate bank-type environment. The main burst of adaptive radiation of the labechiids occurred in the Middle Ordovician, seemingly with the genera derived from a Cystostroma_ or Pseudostylodictyon-like ancestor, as shown in Fig. 10. From the nature of the radiation of skeletonized genera (Figs. 8, 10), it appears that we are viewing a genuine invasion of patch reef and bank habitats previously relatively free from competitors. The early appearance of Stromatocerrzum from the Tasmanian ‘Middle Ordovician’ (Fig. 8), an ‘advanced’ form in terms of its occurrence at the end of an evolutionary pathway (see Stromatocerzum group B in Fig. 10) may merely serve to indicate how rapidly the adaptive radiation took place in the Chazy, rather than to suggest links with a supposed Early Cambrian Stromatocerium archetype (Khalfina & Yavorsky, O7A))e Recognition in some labechiids of immature and mature (or alternating) growth stages — the basal layers of coenostea (and the bases of latilaminae) exhibiting immature stages of growth — may be significant in clarifying phylogenetic relationships within the group (Galloway, 1957). For example, Galloway (1957, p. 394) noted in one specimen of Cystzstroma canadensis the presence of a Rosenella-type ‘Immature’ stage, and Kapp & Stearn (1975) have observed Cystostroma as the ‘Immature’ stage of Labechza prima. Morphologically the most simple, calcified laminar hemispherical forms are Cystostroma, Pseudostylodictyon and Rosenella (Fig. 10). They typically exhibit rows of simple cysts (or laminae in Pseudostylodictyon) arranged in an imbricated manner, denticles on their upper surfaces and mamelons but no pillars. By simple superposition of denticles to form rounded pillars, the genera Labechia (L. prima group), Stratodictyon and Pachystylostroma may be derived. In view of the growth stages exhibited by Labechza (Yavorsky, 1961; Kapp & Stearn, 1975) Cystostroma, rather than Pseudostylodictyon, should be regarded as its ancestor. Stratodzctyon appears to have been derived from a member of the L. prema group, or less likely, directly from Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 116 THE ORDOVICIAN STROMATOPOROIDS (group B ) (+ S. canadense group) (groups A andC) Biede-likerenlars Blade-like Angular pillars & Round-angular pillars Blade-like pillars EN & denticles pillars no denticles & denticles no denticles ‘) : Aw Cry ptophragmus soe ee (Cladophragmus & See Labechiella i 9 Thamnobeatricea) ? Pillars & denticles Solid tissue fills area of pillars & ( | Rosenella | | Ludictyon | é é Pseudostylodictyon Rosenella Ludi udictyon interspaces ) (Plumatalinia) é Denticles Labechia only (Forolinia) Cystostroma Sinodictyon Dermatostroma ANE : Mamelons, Round pillars & pillars & Mamelons & no denticles aeanivales denticles Non-calcified Pillars & denticles ancestor with simple cysts ENCRUSTING LAMINAR — HEMISPHERICAL CYLINDRICAL Fig. 10. Diagram showing possible evolutionary pathways for the Ordovician genera of labechiid stromatoporoids. Pseudostylodictyon (Webby, 1969, p. 653), while Pachystylostroma appears to have evolved, as Kapp & Stearn (1975) have outlined, from Pseudostylodictyon. The culmination of the trend towards long and continuous pillars derived from superposed denticles leads to representatives of Labechza (L. conferta group) with cysts and Labechiella with laminae. Stromatocertum (S. rugosum group) is derived from Labechiella and has irregular (secondarily altered) pillars. None of these latter forms exhibits denticles. Derivation of Lophiostroma and Dermatostroma is more problematical. The clue to the derivation of Lophzostroma with its poorly differentiated internal structures owing to a completely solid infill of tissue is seen in the Middle Ordovician species L. shantungensis which shows traces of original pillars and a few large unfilled cysts in the interspaces, as in a Labechiella or Labechza. Furthermore the upper surface of the coenosteum of the respective type species of Lophzostroma and Labechza have an identical papillose appearance. The encrusting genus Dermatostroma includes forms with solidly fused vertically aligned prisms which may be allied to Lophiostroma (Dermatostroma A in Fig. 10), and forms with a few laminae and pillars which may be related to Labechiella (Dermatostroma B in Fig. 10). Cystistroma, as previously suggested by Webby (1969, p. 653), may have been derived from Pseudostylodictyon with the large, rounded-angular pillars evolving from mamelons, not from denticles. The new Stromatocerium-like genus (Webby, in press a) bears similarities to Cystzstroma in exhibiting denticles on upper surfaces of cysts in the interspaces between the large pillars but differs in having ‘composite’ vane- like pillars, each apparently developed at the site of a former mamelon. It seems to have a different derivation from Cystzstroma, probably from a Pachystylostroma. Other forms with large ‘hollow’ wall-like radiating pillars of S. rugosum type (group A of Fig. 10) and slender, intermeshed, ‘composite’ blade-like pillars, referred to the Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 B. D. WEBBY 117 Stromatocertum michiganense group (group C of Fig. 10) may have a similar origin, but they lack denticles. The cylindrical labechiids appear to have evolved from the _ laminar- hemispherical forms by the development of extended mamelon-like upgrowths perhaps from a Cystostroma-like base (Galloway & St. Jean, 1961). The relationships have not been clearly established but there would appear to be two main trends (Fig. 10) — one through the Chinese fasciculate-cylindrical genera Ludzctyon and Sinodictyon to the large unbranched Aulacera, and the other through the slender North American forms of Cryptophragmus, Cladophragmus and Thamnobeatricea to Alleynodictyon, a genus with blade-like pillars (Webby, 1971). Both lines would appear to commence with a simple Rosenella or Cystostroma-like ancestor. Clathrodictyids may have been derived from a simple labechiid ancestor, but evolved pillars from downward inflections of laminae rather than by superposition of denticles as in labechiids, and developed walled astrorhizae. Adequate time seems to have been available for these morphological changes to have taken place, given a line of descent from a Chazy, Cystostroma-like ancestor to the first appearance of clathrodictyids in the “Trenton’ (Fig. 8). The alternative is a much earlier, independent ancestry for the group, possibly from the Early Cambrian Clathrodzctyon of Vlasov (1961) and Khalfina & Yavorsky (1967). To be directly ancestral to Middle Ordovician and later representatives of the genus would imply that the supposed Early Cambrian Clathrodictyon lost its ability to preserve a mineralized skeleton for a period of 110 m.y. to the Middle Ordovician. It is more difficult to derive the complex morphology of the cliefdenellids from a labechiid. The group may have evolved from a clathrodictyid like Plexodictyon but there would seem to be too limited a period of time within the ‘Trenton’ (Fig. 8) to make all the necessary morphological changes, viz., addition of denticles to the upper surface of its primary laminae, large tube-like pillars and complex astrorhizae. A more realistic view allowing for the development of the complex morphological features is to suggest that cliefdenellids arose as an independent Ordovician group from an earlier, possibly Cambrian soft-bodied ancestor. The marked morphological differences between the three separate families of Ordovician stromatoporoids favour a much earlier origin possibly from a common (? soft-bodied) ancestor in the Cambrian (very doubtfully from within the irregular archaeocyathans, suborder Archaeosyconina), with independent lines of descent through the Ordovician. The initial Middle Ordovician record of each family is based on the appearance of its first skeletonized remains, and does not necessarily coincide with the origins of the individual group. ACKNOWLEDGEMENTS In earlier stages of preparation of this review I was assisted by funds from the Australian Research Grants Committee (A.R.G.C. grant No. E73/15102), and a Royal Society and Nuffield Foundation Commonwealth Bursary. Many of the illustrated thin sections were made by Mr D. G. Morris. References ALLOITEAU, J., 1952. — Classe des Hydrozoaires. In Piveteau, J. (ed.), Trazté de Paléontologie, 1: 377-398. Masson & Cie, Paris. Banks, M. R., 1962.— Ordovician System. In Spry, A. & Banks, M. R. (eds). The geology of Tasmania. /. geol. Soc. Aust., 9: 147-176. Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 118 THE ORDOVICIAN STROMATOPOROIDS BILLINGS, E., 1857.— Ordovician and Silurian rocks of North America. Geol. Surv. Canada, Rept. Progress, 1853-1856: 343-345. BIRKHEAD, P. 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Inst., n.s., 87: 1-160. ZHURAVLEVA, I. T., 1955.— Arkheotsiaty nizhnego kembriya vostochnogo sklona Kuznetskogo Alatau. Trudy Paleont. Inst. Akad. nauk. SSSR, 56: 5-56. ——, 1960. — Arkheotszaty siberskoy platformy: 1-344. Akad. nauk SSSR, Moskva. ——, 1970.— Porifera, Sphinctozoa, Archaeocyathi — their connections. In Fry, W. G. (ed.). The Biology of the Porifera. Symp. zool. Soc. Lond., 25: 41-59. ——, and Miackova, E. I., 1974.— Stavnitelenaya kharakteristika Archaeata 1 Stromatoporoidea. In Sokolov, B.S. (ed.). Drevnzye Cnidaria, 1: 63-70. Nauka, Novosibirsk. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 Freshwater Sponges from the Northern Territory (Porifera: Spongillidae) J. STANISIC STANISIC, J., Freshwater sponges from the Northern Territory (Porifera: Spongillidae) . Proc. Lenn. Soc. N.S.W. 103 (2), (1978) 1979: 123-130. Three spongillids are recorded from a freshwater billabong in Arnhem Land, Northern Territory, Australia. Metania ovogemmata sp. nov. and Radzospongilla streptasteriformis sp. nov. are figured and described. M. ovogemmata represents the first record of the genus Metania Gray in Australia and its discovery establishes a connection between the Australian and South American spongillid faunas. R. streptasterzformzs extends the range of the genus Radzospongilla Penney and Racek in Australia and a study of its relationship to other members of the genus indicates that there has been a radiation of radiospongillids in Australia. Eunapius sinensis (Annandale), previously recorded from the eastern states of Australia, is recorded for the first time from the Northern Territory. This new information forms the basis for further speculation on the origins and relationships of Australian spongillids. J. Stanisic, Department of Malacology, The Australian Museum, P.O. Box A285, Sydney South, Australia 2000; manuscript recezved 16 November 1977, in revised form 18 September 1978. INTRODUCTION Following the comprehensive revision of a worldwide collection of freshwater sponges by Penney and Racek (1968), Racek (1969) completely revised the freshwater sponges of Australia and produced keys to all the described species. However, Racek’s conclusions regarding the origin, dispersal and distribution of the Australian spongillid fauna were based on only limited material from the remote areas of central and northern Australia. Apart from the single record of Radzospongilla hispidula Racek from Darwin and two specimens of Radiospongilla philippinensis (Annandale) from Derby (W.A.), the material treated by Racek (1969) came from localities in the eastern states. The paucity of material from these remote areas is largely due to their inaccessibility to previous collectors. However, with the establishment of the Crocodile Research Facility at Maningrida in Arnhem Land (N.T.), the author was able to obtain a collection of spongillids from Australia’s far north. Detailed taxonomic studies of this collection revealed the presence of two new species of spongillids and the first record of the genus Metanza Gray from Australia. The sponges described in this paper provide new information on the distribution of the Australian spongillid fauna. More importantly however, this new information enables a reassessment of the possible origin of some of Australia’s freshwater sponges to be made. The taxonomic and morphological terminology used here follows Penney and Racek (1968). Preparation of the material followed the outlines presented by Gee CBI) Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 124 FRESHWATER SPONGES TAXONOMY Genus Metania Gray Metanza Gray, 1867, p. 551 Metanza Penney and Racek, 1968, p. 147 (and synonymy) Type species: Spongilla reticulata Bowerbank (1863) This genus, as redefined by Penney and Racek (1968), includes those species which possess a single layer of radially arranged tubelliform gemmoscleres and free microscleres. The genus is restricted to tropical areas of Asia, Africa and South America. The new species, which is described below is the first representative recorded from Australia. Metanza ovogemmata sp. nov. Material: Freshwater billabong (‘Benamanarka Gunora’) near Maningrida, Arnhem Land, Northern Territory, 12°00'S, 134°20'E, coll. G. Wells, 6. x. 76, holotype (Australian Museum Z3693) . Description: Sponge forming encrustations of variable thickness (1-3 mm) on logs in shallow water; surface reticulate and markedly hispid with spicules projecting through the dermal membrane. Skeleton consisting of tracts of spicules which form vague triangular meshes. Oscula conspicuous. Consistency of dry sponge firm. Megascleres of two distinct size classes. The larger series forms the primary skeleton of the sponge, while the smaller series is mainly restricted to the vicinity of the gemmules, forming a spicular envelope around each gemmule. The larger series (Fig. 1 a-c) consists of stout fusiform amphioxea; slightly curved, ranging from smooth to incipiently spined except on the distal portions of the sclere. Length range 220-290 yum, width range 10-20 um. Those in the smaller series (Fig. 1d,e) are short, stout, hastate amphioxea; curved and covered with numerous conical spines which tend to form whorl-like aggregations at the scleral apices. Length range 195-230 um, width - range 12-20 um. Microscleres (Fig. lf,g) slender, fusiform amphioxea; straight to slightly curved and covered with numerous spines. In the central portion of the sclere these spines become longer and complex. Length range 60-90 um width range 2-4 um. Gemmoscleres (Fig. 1h-1) tubelliform with an almost circular rotule at one end and terminating in a smooth knob at the other. The face of the rotule is clear of any striations while the margins are strongly recurved. The shaft of the sclere possesses a number of large conical spines. Length range 25-35 um, width of shaft 2-4 um, diameter of rotule 16-21 um, width of terminal knob 3-7 um. Gemmules (Fig. 1m,n) large and oviform; abundant and scattered throughout the skeletal meshwork. The pneumatic layer of the gemmule is thin with the gemmoscleres embedded radially so that the terminal knob is to the outside. The knob does not project beyond the outer gemmular membrane. Surface of the gemmule is granular. The foramen of the gemmule is tubular and the porus tube projects a short distance beyond the outer gemmular membrane. The gemmules are surrounded by an envelope of spiny megascleres. Dimensions of gemmules 420-520 um (long axis) by 260-370 um (short axis) . Type: Holotype with slides and vial of gemmules in the Australian Museum. Distribution: Hitherto known only from the type locality in Arnhem Land, Northern Territory. Colour: Spirit specimen black. Discussion: The discovery of this new species in tropical northern Australia fills an important gap in the distribution of the genus Metanza. Although known to occur in Proc. LInn. Soc. N.S.W., 103 (2), (1978) 1979 J. STANISIC 125 tropical areas, the absence of material from Australia puzzled Penney and Racek (1968), who concluded that the genus had a discontinuous distribution. It now appears that Metanza, like other groups of spongillids, has a continuous southern distribution. The presence of two classes of megascleres and the peculiar arrangement of the smaller series around the gemmule, suggest that this species is most closely related to Metanza reticulata (Bowerbank) which occurs in the Amazon River of South Cc 3 b S a 3 | h i j f 9 k l 8 : OR n Fig. 1. Metania ovogemmata sp. nov. a-c. larger series of megascleres d-e. smaller series of megascleres k-]. rotules of gemmoscleres f-¢. microscleres m. gemmule h-j. gemmoscleres n. micropyle apparatus Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 126 FRESHWATER SPONGES America. However, the difference in the size of the megascleres of the two species and the fact that these are amphioxea in M. ovogemmata as opposed to amphistrongyla in M. reticulata, indicate that M. ovogemmata deserves separate specific status. The striking arrangement of the hastate amphioxea in M. ovogemmata around the gemmule, appears to be an adaptation indicative of species which have small gemmoscleres and an associated small pneumatic layer in the gemmule. Similar capsules are found in Trochospongilla Vejdovsky, Uruguaya Carter and Drulia Gray, all of which have small radially arranged gemmoscleres and small pneumatic layers. Considering the presence of the two congeners, Metanza vesparia (von Martens) and Metanza vesparioides (Annandale) in the Asian region, the relationship of M. ovogemmata to a South American species has important implications regarding the possible origin of this genus. These implications will be discussed later. Genus Radiospongilla Penney and Racek Radiospongilla Penney and Racek, 1968, p. 61 (and synonymy). Radiospongilla Racek, 1969, p. 279. Type species: Spongilla sceptroides Haswell (1882) . Penney and Racek (1968) introduced this genus as a link between those spongillids with acerate gemmoscleres and those with birotulate gemmoscleres, showing that the previous grouping of the spongillids into the two sub-families Spongillinae and Meyeninae was unwarranted. Hitherto six species are known from Australia. Radtospongilla streptastertformis sp. nov. Materzal: Freshwater billabong (‘Benamanarka Gunora’) near Maningrida, Arnhem Land, Northern Territory, 12°00'S, 134°20’E, coll. G. Wells, 6. x. 76, holotype (Australian Museum Z3695) . Description: Sponge forming very thin encrustations in the corrugations of bark from logs in shallow water. Surface of the sponge is an irregular network of spicules with some of these projecting through a poorly developed dermal membrane giving the sponge a slightly hispid appearance. Megascleres (Fig. 2a,b) slender to stout, curved fusiform amphioxea; strongly spined over the entire length with no distinct aggregation of the spines. Length range 160-200 um, width range 4-10 pm. wrvool a b Fig. 2. Radzospongilla streptasteriformis sp. nov. a-b. megascleres c-f. gemmoscleres g. gemmule Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 J. STANISIC 127 Microscleres absent. Gemmoscleres (Fig. 2c-f) short, stout abrupt amphistrongyla covered with numerous long straight spines. Occasionally the central portion of the sclere is less spined than the distal portions. Straight to slightly curved. Length range 40-65 um, width range 4-6 ym. Gemmules (Fig. 2g) abundant and scattered throughout the symplasm of the sponge. These are large and spherical with a well developed pneumatic layer. The gemmoscleres are embedded radially in the pneumatic layer and project beyond the outer gemmular membrane, making the surface of the gemmule distinctly hispid. The porus tube does not project beyond the outer gemmular membrane and is surrounded by a conical depression. Diameter of gemmule 340-440 um. Type: Holotype with slides and vial of gemmules in the Australian Museum. Distribution: Hitherto known only from the type locality in Arnhem Land, Northern Territory. Colour: Not reliably recorded. Discussion: The discovery of Radzospongiulla streptasteriformis in the alkaline waters of Australia’s north, suggests that the genus Radzospongilla Penney and Racek may not be as restricted in distribution as Racek (1969) indicated. Based largely on collections from the eastern seaboard, Racek suggested that most species of this genus preferred acidic waters. The gemmoscleres of R. streptasteriformzs which are small and evenly spined abrupt amphistrongyla, readily separate this species from its congeners. However, the dimensions of the megascleres and gemmoscleres indicate that this species is closely related to Radzospongilla synoica Racek and Radzospongilla cantonensis (Gee) . These three spongillids are characterized within the radiospongillids by their small megascleres and small gemmoscleres. They show a range of gemmosclere structure from amphioxea (R. synozca) through abrupt amphistrongyla (R. streptasteriformis) to the formation of almost perfect pseudorotules (R. cantonensis) . As R. synotca is endemic to eastern Australia and R. cantonenszs is also present in eastern Australia (unpublished data), the distribution of these three species indicates a localized radiation from the central stem of radiospongillids as represented by the extant species Radzospongilla cerebellata (Bowerbank) and Radzospongilla philippinensis (Annandale). The evolutionary importance of the genus Radiospongilla (Penney and Racek, 1968; Racek, 1969; Racek and Harrison, 1975), makes the occurrence of such a radiation in the Australian region assume particular relevance in discussions concerning the origins of spongillids. The significance of this point will be discussed later. Genus Eunapzius Gray Eunapius Gray, 1867, p. 552 Eunapius Penney and Racek, 1968, p. 21 (and synonymy). Eunapius Racek, 1969, p. 271 Type species: Spongilla cartert Bowerbank (1863) This genus is characterized by species which have a tangential arrangement of gemmoscleres around the gemmule and which lack free microscleres. Members of the genus are widely distributed and four species are recorded from Australia. Eunapius sinensis (Annandale) is known from the eastern and western river systems in New South Wales and Queensland. It is here recorded for the first time from the Northern Territory. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 128 FRESHWATER SPONGES Eunaptus stnensis (Annandale) Spongilla (Stratospongilla) sinensis Annandale, 1910, p. 183. Eunapius sinensis Penney and Racek, 1968, p. 35 (and synonymy). Eunapius sinensis Racek, 1969, p. 273. Materzal: Freshwater billabong (‘Benamanarka Gunora’) near Maningrida, Arnhem Land, Northern Territory, 12°00’S, 134°20’E, coll. G. Wells, 6. x. 76 (AM Z3695). Type: Of S. senenszs in the collection of the United States National Museum. Distributzon: According to Penney and Racek (1968), ranging from USSR through Manchuria and mainland China to Australia. Not known from SE Asia. Discussion: The characteristics of this specimen in both its mode of growth and spicular components make it fully comparable with the previously recorded Australian specimens. Racek (1969) considered that the thick pneumatic coat of the gemmule, which is reinforced by tangential layers of robust gemmoscleres, makes this species particularly suited to harsh environments. The specimen was found growing in very close association with M. ovogemmata. However, the pavement layers of gemmules of E. s¢nenszs were readily discernible from the more loosely scattered gemmules of M. ovogemmata. GENERAL DISCUSSION Origins and Relationships of the Australian Spongillids Racek (1969) believed that most of the Australian spongillids were a southern extension or represented mere races of the Asian fauna. He based this assessment on the distribution of the extant spongillids in these two areas. More recently however, Racek and Harrison (1975) have reconsidered this theory. Racek and Harrison examined the phylogenetic position of the fossil spongillid Palaeospongilla chubutensss Ott and Volkheimer which was discovered in the lacustrine sediments of the Cretaceous of Argentina. Consideration was given to the southern distribution of a number of the extant spongillids which showed close affinity with the fossil. In particular, Racek and Harrison referred to the presence in Australia of members of the genus Radzospongilla Penney and Racek, which could be related to the Cretaceous fossil, and the distribution of Spongzlla alba Carter in both Australia and South America. These workers considered the pattern of continental drift in an attempt to reconstruct dispersal routes leading from the Cretaceous fossil to the present day distribution of spongillid fauna. Racek and Harrison concluded that it was more reasonable to consider that Asia and Australia had independent faunal gains along dispersal routes which existed prior to the dismemberment of the great southern continent Gondwanaland. Two dispersal routes leading west-east were suggested. One route was along the connected South American, Antarctic and Australian plates, while the other was through Africa and Indo-Pakistan. Subsequently there may have been interchange between the Asian and Australian faunas. However, the evidence for these conclusions was based almost solely on the assumption that the extant fauna was directly related to the Cretaceous fossil. With the discovery of Metanza ovogemmata sp. nov. in Australia, a reassessment of spongillid dispersal routes is possible. It has already been demonstrated that M. ovogemmata is more closely related to Metanza reticulata (Bowerbank) of South America than to its Asian congeners. The affinity between these extant spongillids can only be reasonably explained by divergence from a common stock. In order to account for the present distribution of these two species, subsequent dispersal would have required connected land masses which were present during the Mesozoic when Australia and South America were part of Gondwanaland (Smith, 1971). Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 J. STANISIC 129 Although Racek and Harrison (1975) came to similar conclusions regarding the importance of Gondwanaland in spongillid dispersal they considered only dispersal routes which were west-east along the Mesozoic continuity. A study of the distribution of the genus Metanza Gray reveals the possibility of dispersal in an east-west direction. Metanza vesparia (von Martens) and Metanza vesparioides (Annandale) which occur through Africa, Burma, Borneo and Indonesia indicate dispersal and speciation from South America to Africa and Indo-Pakistan. As M. ovogemmata possesses a number of ancestral traits, e.g. two classes of amphioxeous megascleres, it is possible to demonstrate an evolutionary series from M. ovogemmata through M. retzculata to the Asian congeners. Therefore, in this case dispersal and speciation has most probably occurred in an east-west direction with the genus Metanza having its origins in the Australian region. Although little is known of the origin of the spongillids with birotulate gemmoscleres (e.g. Metanza) , they are considered to have arisen from an hypothetical stock from which the extant genus Radzospongilla Penney and Racek has also been derived (Penney and Racek, 1968). The gemmoscleres of Radzospongilla cantonensis (Gee) and Radzospongilla crateriformzs (Potts) possess almost perfect rotules, suggesting that these two species share a common ancestry with the birotulate genera. In order to demonstrate more conclusively the emergence of birotulate genera in Australia, it is necessary to consider the distribution of radiospongillids which are considered to be phylogenetically related to these genera. However, this requires a reassessment of the suggested relationships within the genus Radzospongilla. Penney and Racek (1968) considered that R. cantonensis and R. crateriformis form part of an evolutionary line within the radiospongillids. Their conclusions were based on the fact that the gemmoscleres of both these species possessed pseudorotules. This view implies that the birotulate genera have had a monophyletic origin. Such a view, however, does not explain the marked morphological diversity of the birotulate genera. Moreover, this view leads to difficulties in explaining relationships between the distribution of these radiospongillids and the distribution of the birotulate genera. Recent developments based on the work of Poirrier (1974, 1976) and Stanisic (1977) enable an alternative view to be put forward. These workers have shown that using gemmoscleral form as a first principle for establishing relationships within genera, is not entirely valid. Such features are subject to ecomorphic variation which can mask true relationships. If, on the other hand one considers the length of the megasclere as a basis for assessing relationships, a new and more reasonable argument presents itself. The difference in the lengths of the megascleres of R. cantonensis and R. crateriformis indicates that these two species have diverged from the central line of the radiospongillids at different times. The central line is represented by the Radiospongilla cerebellata (Bowerbank) group. Hence it is possible that the birotulate genera have arisen from more than one ancestor and that their origins are polyphyletic. Using megasclere length as a criterion, it is now also reasonable to suggest that R. cantonenszs is more closely related to Radzospongilla synoica Racek and Radvospongilla streptasteriformis sp. nov. as all three spongillids have small megascleres. In view of the fact that these three spongillids occur almost solely in the Australian region, it is probable that they represent a radiation from a common ancestor in this region. Therefore, if R. cantonensts shares a common ancestry with some of the birotulate genera in Australia it is possible to document the emergence of birotulate genera in Australia. This not only supports the views of the present study regarding the origins of the genus Metanza but also the view of Racek (1969), who considered that Heterorotula Penney and Racek evolved in Australia. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 130 FRESHWATER SPONGES Inter-generic relationships within the family Spongillidae still need to be reliably established. The broad hypothesis regarding the relationship of the genus Radiospongilla to the birotulate genera which was put forward by Penney and Racek (1968), while providing some insight, is too vague to provide the explanatory and predictive powers which would make such a hypothesis useful in determining relationships. On the other hand, the specific arguments which have been presented above, can be used to demonstrate the emergence of birotulate genera in Australia. It is now also possible to predict a relationship between the radiospongillids mentioned above and these birotulate genera. Although highly speculative, such a hypothesis is favoured by the fact that its restrictive nature makes it open to testing by future research and discoveries. As mentioned at the beginning of the discussion, the origins and affinities of the Australian freshwater sponges have been discussed by Racek (1969) and Racek and Harrison (1975). The new material documented in the present study, has enabled a further assessment to be made of the considerations initiated by these workers and it is now possible to show a definite connection with the unique South American spongillid fauna. Reconsideration of inter-generic relationships and distribution patterns indicate the possibility of an Australian origin for some of the birotulate genera, in particular Metanza. The information provided by this new material however, emphasizes the need for further collecting within the Australian region. ACKNOWLEDGEMENTS I am grateful to Mr. Graeme Wells of the Crocodile Research Facility, for obtaining this valuable material. I would also like to thank Patsy Armati-Gulson, Dr. Allin Hodson and Associate Professor John Simons of the School of Biological Sciences, University of Sydney, for their time and constructive criticism during the drafting of the manuscript. Thanks are also due to Miss Bethelle Heuer for her care in the preparation of the typescript. References ANNANDALE, N., 1908.— Notes on freshwater sponges. IX. Preliminary notice of a collection from Burma with the description of a new species of Tubella. Rec. Indian. Mus., 2: 157-158. BOwWERBANK, J. S., 1863. — A monograph of the Spongillidae. Proc. Zool. Soc. Lond., 1863: 440-472. Gre, N. G., 1931.— The study of freshwater sponges as a hobby. Vzct. Nat., 48: 4-7. Gray, J. E., 1867.— Notes on the arrangement of sponges with the description of some new genera. Proc. Zool. Soc. Lond., 1867: 492-558. PENNEY, J. T., and Racek, A. A., 1968. — Comprehensive revision of a worldwide collection of freshwater sponges (Porifera: Spongillidae). U.S. Nat. Mus. Bull., 272: 1-184. PoIRRIER, M. A., 1974.— Ecomorphic variation in gemmoscleres of Ephydatza fluviatilis (Linnaeus) (Porifera: Spongillidae) with comments upon its systematics and ecology. Hydrobiologia, 44: 337- 347. —____, 1976. — A taxonomic study of the Spongzlla alba, S. cenota, S. wagneri species group (Porifera: Spongillidae) with ecological observations of S. alba. In: “Aspects of Sponge Biology.” F. W. Harrison and R. R. Cowbsn, eds, Academic Press, N.Y: 203-213. Racek, A. A., 1969.— The freshwater sponges of Australia. (Porifera: Spongillidae). Aust. J. mar. Freshwat. Res., 20: 267-310. , and Harrison, F. W., 1975.— The systematic and phylogenetic position of Palaeospongzlla chubutensts (Porifera: Spongillidae). Proc. Linn. Soc. N.S.W., 99: 157-165. SMITH, A. G., 1971.— Continental Drift. In: Gass, I. G., Smiru, P. J., and Witson, R. C. L., “Understanding the Earth”. Horsham, Sussex: Artemis Press: 213-231. STANISIC, J., 1977.— Studies on the freshwater sponge Radzospongilla sceptrotdes (Haswell, 1882) (Porifera: Spongillidae) . Univ. Sydney, unpubl. M.Sc. thesis. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 Hans Laurits Jensen, 1898-1977 MACLEAY BACTERIOLOGIST, 1929-1947 (Memorzal Serzes No. 26) Hans Laurits Jensen was born 27 June 1898 at Frederiksund, Denmark, and died in Copenhagen on 5 October 1977. He came to Australia in 1929 to take up the position of Macleay Bacteriologist of the Society, a post which he occupied for the next eighteen years until his return to the State Plant Laboratory at Lyngby in Denmark. Before coming here he trained in agronomy at his home university (the Royal Danish Veterinary and Agricultural High School), after which he had been appointed Bacteriologist in the State Plant Laboratory. A Rockefeller Grant then enabled him to work for two years with H. G. Thornton at the Rothamsted Experimental Station before his Linnean Society post in which he was to make such a notable contribution. Work on free-living nitrogen fixation which largely occupied the middle part of his Australian period earned Jensen the doctorate of his university. From 1943 to 1947 he combined his continuing research programme with lecturing in Bacteriology to science students at the University of Sydney. Returning to Denmark in 1947, he was in charge of bacteriology in the State Plant Laboratory, completing his service with seven years as Director until his retirement in 1968. During his career Dr Jensen published more than 170 articles, of which 35 were very substantial contributions to the Proceedings of this Society. He became recognized as an outstanding agricultural microbiologist, particularly in connection with organic matter decomposition, nitrogen fixation and soil fertility; he was the recipient of many awards, both in his native Denmark and on the world scene. Hans Jensen played a significant editorial role in several internationally recognized journals and was commonly consulted as examiner and research adviser. It was a measure of the man’s influence, and of the high regard in which he was held, that colleagues and friends commemorated his seventieth birthday with a volume of scientific articles by thirteen contributors from eight countries representative of north-south and east-west hemispheres. * Dr Jensen’s research interests were wide indeed, ranging from several major contributions to the activity, nature and systematics of soil bacteria (notably actinomycetes and corynebacteria) to more immediately practical matters concerned with the decomposition of soil organic matter, the degrading of diverse herbicides and other potent organic molecules which under present day farming practice find their way into the soil. At no stage did Jensen restrict himself to a narrow field although, of course, the emphasis was adjusted from time to time. For example his interest in biological nitrogen fixation was maintained over some thirty years and included many investigations with both free-living and symbiotic forms. Particularly it was Jensen’s critical quantitative studies of the significance and requirements of non-symbiotic nitrogen fixation which brought about a more realistic re-assessment of its limited contribution to maintaining soil nitrogen under conditions of agricultural practice in Australian wheat production. At the same time Jensen contributed most significantly to a better knowledge of the taxonomy of the free-living aerobic nitrogen-fixing bacteria as well as the pasture legume: Rhzzobzum association. Study of the latter was initiated during the last part of his period as Macleay Bacteriologist, at a time when *Festskrift til Hans Laurits Jensen — Gadgaard Nielsens Bogtrykkeri, Lemvig, 1968. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 132 HANS LAURITS JENSEN, 1898-1977 there had been very little sound work along these lines accomplished on the Australian scene, but when this country’s work on symbiotic nitrogen fixation was about to take off at a greatly accelerated pace. Jensen’s work in Australia and, less intensively, after his return to Denmark, remains a valuable part of our body of knowledge of this vastly important symbiosis. Besides these major continuing interests Jensen was responsible for comprehensive articles in the wider field of soil microbiology and methodology. According to the needs of the occasion (including the war years) he was also prepared to give his time ungrudgingly to the investigation of practical aspects of microbiology which took in such diverse topics as the suitability of substitute agars, dew-retting of flax, ensilage, antifungal preservatives and the safe storage of blood. For most of his stay in Australia Jensen was of necessity a lone worker, obliged to make the best he could of a quasi-official arrangement between the Society and the University of Sydney. A shrinking effective budget as post-war inflation leapt ahead at the same time as the income from the Macleay bequest remained static, or declined, was a constant restraint. The late Professor Hugh Ward helped by extending the hospitality of the Bacteriology Department to provide accommodation and some basic facilities; considerable support was provided by several Banks, particularly in providing a greenhouse and permitting the employment of a graduate research assistant. Despite these limitations Jensen was able by temperament and ability to use his time in Australia as probably the most productive period of his career. He did this with a series of deceptively simple but beautifully designed and executed experiments that gave clear answers to well directed questions. At times indeed so economical of time and effort was his experimentation, so free his work area of clutter, it seemed that papers could be written directly from the laboratory bench. Hans Jensen truly belonged to the classic tradition of Winogradsky and Beijerinck; his work was a conjunction of impeccable technique and ability to think a problem through to a logical and practical significant conclusion — qualities sometimes missing from contemporary “black-box, publish-or-perish” exponents of the art. We count ourselves privileged to have enjoyed his professional co-operation and personal friendship both in Australia and when we visited him, his wife Helene and his family in Denmark. Agricultural microbiology is in so many ways the better for Hans Jensen’s long and distinguished contributions. Y. T. TCHAN J. M. VINCENT Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 A KEY TO THE MEMORIAL SERIES OF THE LINNEAN SOCIETY OF NEW SOUTH WALES (1928-1978) In October 1978, the Council of the Society resolved to commission an essay on the late Dr Hans Laurits Jensen, Macleay Bacteriologist from 1929 to 1947, for the Memorial Series. The essay, twenty-sixth in the series, is printed in this issue (pp. 131-132). It is the latest in a line now spanning more than fifty years of publication. Council believes members will be interested in the record of that half-century and, accordingly, has ordered the issue of a key to the series. A footnote to the first article, dated 1928, explained the original intention: “The Memorial Series will comprise memorials prepared, from time to time, under the direction of Council, of distinguished Members of the Society who have died’. That has remained a guide to successive Councils. As a result the Society possesses a valuable biographical record, a record believed to be unique among the scientific societies of this country until the Australian Academy of Science began to publish memorials of its deceased Fellows about twenty years ago. Each memorial article was published in the Society’s Proceedings and in limited numbers of separate offprints for private distribution. Until at least the early 1960s separate issues generally appeared with distinctive olive-green or grey printed covers. More recently they have come in humbler style. Of the entries in the following list, only that for J. J. Fletcher calls for particular comment. It alone was not printed as such in the Proceedzngs. In fact, only the separate issue indicates its place in the series. This unusual situation arose because the appreciation by Baldwin Spencer had already appeared in the journal before Council decided to establish the Memorial Series. The Haswell memorial set a pattern and so when Council resolved to commemorate Fletcher in the same style a list of his publications was needed to supplement Spencer’s article. That list, compiled by the then Secretary (A. B. Walkom) was printed in the Proceedings for 1929. The separate issue of the Fletcher memorial (1929) consists of the list of papers appended to reprints of the text and portrait first published two years earlier. Like its immediate neighbours in the series, the Fletcher separate has pagination different from that of copy in the Proceedings. The practice of re-numbering pages for offprints was abandoned after issue of No. 3. THE MEMORIAL SERIES Nos. 1-25 (1928-1978) 1 WILLIAM AITCHESON HASWELL, 1854-1925 (By H. J. C[arter ]) Proceedings 53, 1928: 485-498. Plate. 2 JOSEPH JAMES FLETCHER, 1850-1926 (By W. B[aldwin] S[pencer] (and) [A. B. Walkom |) Proceedings 52, 1927: xxxiii-xliii. Plate. Proceedings 54, 1929: 686-687. 3 JOSEPH HENRY MAIDEN, 1859-1925 (By A. H. S. L[ucas]) Proceedings 55, 1930: 355-370. Plate. 4 RICHARD HIND CAMBAGE, 1859-1928 (By E. C. A[ndrews ]) Proceedings 59, 1934: 435-447. Plate XIX. 5 CHARLES HEDLEY, 1862-1926 (By C. A[nderson ]) Proceedings 61, 1936: 209-220. Plate X. 6 TANNATT WILLIAM EDGEWORTH DAVID, 1858-1934 (By H. J. C[arter] and W. R. B[ rowne]) Proceedings 61, 1936: 341-357. Plate XVII. 7 ARTHUR HENRY SHAKESPEARE LUCAS, 1853-1936 (By H. J. C[ arter |) Proceedings 62, 1937: 243-252. Plate XII. 8 WALTER WILSON FROGGATT, 1858-1937 (By A. B. W[alkom]) Proceedings 67, 1942: 77-81. Plate III. 9 HERBERT JAMES CARTER, 1858-1940 (By A. B. W[alkom]) Proceedings 68, 1943: 91-94. Plate IV. 10 ALEXANDER GREENLAW HAMILTON, 1852-1941 (By E. C. A[ndrews ]) Proceedings 69, 1944: 176-184. Plate III. 11 ROBIN JOHN TILLYARD, 1881-1937 (By J. W. E[ vans ]) Proceedings 71, 1946 (1947) : 252-256. Plate XVII. 12 CARL ADOLPH SUSSMILCH, 1875-1946 (By E. C. Andrews) Proceedings 73, 1948: 242-248. Plate XIV. Proc. LINN. Soc. N.S.W., 103 (2), (1978) 1979 134 13 14 15 16 17 18 19 20 21 22 23 24 ae) KEY TO THE MEMORIAL SERIES ERNEST CLAYTON ANDREWS, 1870-1948 (By W. R. B[rowne] and A. B. W|[alkom ]) Proceedings 77, 1952: 98-103. Plate II. GUSTAVUS ATHOL WATERHOUSE, 1877-1950 (By A. B. W[alkom] and A. J. N [icholson ]) Proceedings 78, 1953 (1954) : 269-275. Plate XVIII. GEORGE DAVENPORT OSBORNE, 1899-1955 (By W. R. B[rowne |) Proceedings 82, 1957: 252-256. Plate XII. WILLIAM NOEL BENSON, 1885-1957 (By W. R. B[rowne]) Proceedings 84, 1959 (1960) : 403-409. Plate XXII. SIR DOUGLAS MAWSON, 1882-1958 (By J. B. C[leland ]) Proceedings 84, 1959 (1960) : 410-414. Plate XXIII. ANTHONY MUSGRAVE, 1895-1959 (By G. P. W[hitley ]) Proceedings 86, 1961: 122-125. Plate V. ‘THEODORE CLEVELAND ROUGHLEY, 1888-1961 (By G. P. W[hitley ]) Proceedings 86, 1961 (1962) : 295-298. Plate XI. ALAN NEVILLE COLEFAX, 1908-1961 (By I. B[ennett }) Proceedings 87, 1962: 220-222. Plate IV. ANTHONY REEVE WOODHILL, 1900-1965 (By D. J. Lee and D. F. Waterhouse) Proceedings 92, 1967 (1968) : 285-297. Plate XVIII. WALTER LAWRY WATERHOUSE, 1887-1969 (By I. A. Watson) Proceedings 95, 1970 (1971) : 260-263. GILBERT PERCY WHITLEY, 1903-1975 (By T. G. Vallance) Proceedings 101, 1976 (1977) : 256-260. WILLIAM ROWAN BROWNE, 1884-1975 (By T. G. Vallance) Proceedings 102, 1977: 76-84. ARTHUR BACHE WALKOM, 1889-1976 (By T. G. Vallance) Proceedings 102, 1977 (1978) : 148-155. T. G. VALLANCE Corrigendum In the W. R. Browne memorial (Proceedings, 102, 1977) on page 80, lines 28-29, the words‘. . . had graduated from the university.’ should have been printed ‘. . . left the university.’, as in the original typescript. Proc. Linn. Soc. N.S.W., 103 (2), (1978) 1979 ADVICE TO AUTHORS The Linnean Society of New South Wales publishes in its Proceedings original papers and review articles dealing with biological and earth science. 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A New Species of Microscolex (Diplotrema) (Annelida: Shee ee from New South Wa 43 A. FAINand R. DOMROW The Family Hypoderidae (Acari) in Australia 47 W.B.K.HOLMES and S. R. ASH An Early Triassic Megafossil Flora from the Lorne Basin, a South Wales ofl. Re CG: RUSSELL, M.C. DEBENHAM and D. J. LEE ' A Natural Habitat of the Insect Pathogenic Fungus pcos in the Sydney ees 75 THELINNEAN SOCIETY OF NEW SOUTH WALES Record of the ANNUAL GENERAL MEETING 1978, Reports and Balance Sheets PART 2 83 B.D. WEBBY : Presidential Address 1978. The Ordovician Stromatoporoids 123 J. STANISIC 7 : Freshwater Sponges from the Northern Territory (Porifera: Spongillidae) 131. MEMORIAL SERIES No. 26 Hans Laurits Jensen, 1898-1977 131 AKEY TO THE MEMORIAL SERIES OF THE LINNEAN SOCIETY OF NEW SOUTH WALES (1928-1978) _ Printed by. Southwood Press Pty Limited, 80-92 Chapel Street, Marrickville 2204 PROCEEDINGS. of the | LINNEAN | SOCIETY NEW SOUTH WALES VOLUME 103 PARTS3&4_ Sa = = Cai ta oe x a Ht hi if “\ NATURAL HISTORY IN ALL ITS BRANCHES THE LINNEAN SOCIETY OF NEW SOUTH WALES Founded 1874. Incorporated 1884. The Society exists to promote ‘the Cultivation and Study of the Science of Natural History in all its Branches’. It holds meetings and field excursions, offers annually a Linnean Macleay Fellowship for research, contributes to the stipend of the Linnean Macleay Lecturer in Microbiology at the University of Sydney, and publishes the Proceedings. Meetings include that for the Sir William Macleay Memorial Lecture, delivered biennially by a person eminent in some branch of Natural Science. The Society’s extensive library is housed at the Science Centre in Sydney. Membership enquiries should be addressed in the first instance to the Secretary. 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VALLANCE — Department of ee & Geophysics, University of Sydney, Australia, 2006. Librarian: PAULINE G. MILLS Linnean Macleay Fellow: JENNIFER M. E. ANDERSON Linnean Macleay Lecturer in Microbiology: K.-Y. CHO Auditors: W. SINCLAIR & Co. The office of the Society is in the Science Centre, 35-43 Clarence Street, Sydney, N.S.W., Australia, 2000. Telephone (02) 290 1612. ©Linnean Society of New South Wales 1 resigned 20 September 1978 ? resigned 23 August 1978 3 appointed 6 September 1978 : * appointed 22 November 1978 1 Cover motif: Clezthrolepis grynulata Egerton Triassic, New South Wales PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 103 PART 3 Fenestrate Bryozoans with Large Apertural Form in the Carboniferous of Eastern Australia BRIAN A. ENGEL ENGEL, B.A. Fenestrate bryozoans with large apertural form in the Carboniferous of eastern Australia. Proc. Linn. Soc. N.S.W. 103 (3), (1978) 1979: 135-170. Study of Australian Carboniferous cryptostome fenestrate bryozoans has revealed that the number of rows of zooecial apertures per branch is transitional between species of Fenestella Lonsdale and Polypora M’Coy and is thus not necessarily the best basis for generic distinction. Apertural studies have demonstrated that simple fenestrate apertures fall into two size distributions with the larger diameter forms bearing a wide, low, peristomal collar in association with an operculum-like covering which bears a central boss-like projection. Species belonging to this group include most Australian Carboniferous forms previously referred to Polypora together with several two-rowed species which, otherwise, would normally be assigned to Fenestella. To separate these large apertural forms, two very closely related genera have been erected, Australopolypora gen. noy. and Australofenestella gen. nov., the former containing the multi-rowed species and the latter the two-rowed species. Six species have been assigned to Australopolypora of which two are new: Aus- tralopolypora rawdonensis and Australopolypora keppelensis parvula. Seven species have been referred to Australofenestella, four of which are new: Australofenestella brookert, Australofenestella trevallynensis, Australofenestella(?) keepitensis, and Australofenestella stroudensts minuta. Brian A. Engel, Department of Geology, University of Newcastle, Newcastle, Australia 2308; manuscript received 10 January 1978, accepted in revised form 26 June 1978. INTRODUCTION Previously described species of fenestrate bryozoans with three or more rows of apertures per branch, from the Carboniferous strata of eastern Australia, have all been placed in Polypora M’Coy (1844) without generic comment, obviously because this broadly defined genus was the only available taxon suitable for their reception. In total, eight such species have been described, five by Crockford (1947, 1949) and three by Campbell (1961). The original diagnosis of Polypora (M’Coy 1844) required these fenestrate forms to have “three to five rows of apertures per branch, apertural peristomes which were never raised and non-poriferous dissepiments”’. These broad generic limitations were not strictly observed even by M’Coy who in the same paper described Polypora verrucosa M’Coy, in which the peristomal collars were strongly elevated (ude Miller 1963). It is thus not surprising that several of the above Australian species, which also exhibit strong peristomal exsertion, were placed by their authors in Polypora. Over the long interval since M’Coy’s diagnosis of Polypora was first published, it has been common practice to place regular fenestrate species, with non-poriferous dissepiments, into either Fenestella or Polypora, depending for the primary generic assignment almost entirely upon the number of rows of zooecial apertures per branch. Effectively, this aspect has assumed a weighting in taxonomy considered to be out of proportion to its importance for reasons given below. The relatively few species which have escaped this rigid dichotomy have depended upon the presence of some exotic Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 136 FENESTRATE BRYOZOANS structural aspect, as can be found in various contemporary genera such as Archimedes Owen 1838, Hemitrypa Phillips 1841 and Semzcoscinium Prout 1859. Because of the virtually uncontrolled addition of many hundreds of species to Fenestella and Polypora, both broadly-defined genera have become totally unwieldy, impractical units for which only tentative efforts have been made in more recent times to devise some scheme of subdivision. With respect to the genus Polypora, the only major attempt at a key classification has been made by Russian palaeontologists who, as a result of their experience over some forty years, have based their grouping of species largely upon the number of rows of zooecia per branch, thus extending the style of demarcation between Fenestella and Polypora to the various species of Polypora with higher numbers of zooecial rows. Australian experience with a limited Carboniferous-Permian sample would suggest that this method can have direct stratigraphic relevance but at the same time it obscures some of the more basic morphological and genetic relationships which may lead to a more satisfactory subdivision. Revised diagnosis of POLYPORA M’Coy. Appreciating the problems accompanying the unrestricted growth of Polypora, Miller (1963) re-described M’Coy’s type material with the object of establishing a suitable standard for subsequent investigations. This emended generic diagnosis appears to have been written largely with the morphology of the type species in mind. In this respect it is rather restrictive in some morphological aspects, variations of which may be considered legitimately to fall within the limits of the genus. In particular, apart from the regular multi-rowed fenestrate features, Polypora is additionally re-defined as having (a) non-carinate branches; (b) regularly spaced, elevated nodes situated on the centreline of the branch; (c) hexagonal-polygonal zooecial bases arranged side by side on a flat basal plate; (d) a microstructure of tubules in the branch wall surrounding apertural margins and forming cores of nodes. Australian Carboniferous species do not readily conform with some of these requirements for the following reasons. Nodes, if developed at all, are scattered in a relatively haphazard fashion over the branch surface being placed adjacent to the proximo-central rim of an aperture in any row. There is a tendency to approach a vaguely central arrangement in a number of the species but none of them has “regularly spaced, elevated nodes situated on the centreline of the branch”, the lack of which, given the current assessment of nodal function, is not considered to be sufficient reason for their exclusion from the genus. In addition, the Australian material is preserved only as external and internal moulds with an absence of primary skeletal remains and hence the inclusion of tubular microstructure as an essential aspect becornes an impractical requirement. It is perhaps relevant to comment that an apparent lack of data on both nodes and micro- structure in Polypora marginata M’Coy did not deter Miller (1963) from regarding it as a legitimate member of the revised genus. As a possible alternative to the key classification of the Russian school, Miller suggested that further subdivision of Polypora could ultimately be based upon the following features: (a) presence or absence of nodes on the obverse of the branches; (b) arrangement of these nodes with respect to a central carina; (c) organization and location of the zooecial chambers in relationship to the ‘basal plate’. Apparently this suggested scheme was intended to result in new genera/sub- genera, for the absence of nodes and the presence of a median carina would appear to be incompatible with his emended diagnosis of Polypora. Over the decade since these suggestions were put forward, ideas on the importance of nodes have changed, a situation foreshadowed by Miller (1963, p.168). Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 137 Personal experience with Australian fenestrates appears to downgrade nodal distribu- tion to the species or even varietal level, and it is difficult to justify any greater impor- tance in taxonomy. Tavener-Smith (1973) has also questioned the validity of Miller’s revised diagnosis regarding nodal occurrence and has rejected the requirements as unwarranted and over-restrictive. MORPHOLOGICAL CONSIDERATIONS Having questioned the conventional relationship between Fenestella and Polypora and finding little else in the definition of either genus that is convincingly diagnostic, a further search was made for other features which could be of some value. An intensive morphological and statistical treatment was made of approximately 300 specimens including all type specimens of Australian Carboniferous species previously referred to either Fenestella or Polypora. This study has demonstrated predictably that much more attention should be given to individual zooecial characteristics. Whilst inter- and intra-zooecial variation is obviously basic in all descriptions, zoarial details have tended to dominate taxonomic studies at the generic/specific level almost to the exclusion of individual zooecial details. Historically, it has been easier, largely because of indifferent preservation, to record such items as zooecial spacing rather than to delineate the characteristics of individual chambers. In more recent years, most authors have realized the need to include zooecial characteristics and such items as the basal outline of zooecial chambers now form a common component of most new descriptions. The following observations on morphological details have arisen from the present study : Number of zooecial rows per branch. The conventional method of subdivision of species between Fenestella and Polypora on the basis of either two or three (and more) zooecial rows per branch was found to be increasingly impractical for Australian Car- boniferous fenestrates. There exists a trend for two-rowed species, commonly assigned to Fenestella, to develop a third row of apertures prior to each branch bifurcation throughout the Carboniferous interval. Low zonal species have no pre-bifurcation apertures, mid-zonal species normally have one or two additional pre-bifurcation apertures and by the late Carboniferous there are commonly at least five or six and often more apertures arranged in a very distinct third row which extends for a con- siderable distance back towards the previous branch division. Since dominantly three- rowed species, normally placed in Polypora, commonly have a reduction to two rows of apertures for some distance immediately post-bifurcation, it is readily apparent that these tendencies lead to species which are obviously neither two- nor three-rowed in character (Fig. 3, 9). This problem of generic separation has already been cited by Campbell (1961) in the case of Fenestella? alttnodosa Campbell where after doubtfully assigning the species to Fenestella, this author concluded that “perhaps it would be equally well placed in Polypora” (Campbell, 1961, p.459), a change which has now been proposed herein for other morphological reasons. Many previous authors have noted the undoubtedly polyphyletic nature of both Fenestella and Polypora but none appears to have questioned the validity of using the number of apertural rows as a primary generic indicator. Based upon the present study of Australian species, it is suggested that at least some of the present taxonomic difficulties stem from this assumption. The change from two to three rows of apertures would appear to be a transitional one which does not seem to have been discussed previously with sufficient clarity. Excellent support for this transitional change comes from the genus Septatopora (Engel, 1975), where other highly diagnostic morpholo- gical features result in this one genus containing a group of species which progressively Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 138 FENESTRATE BRYOZOANS change from two to four or five zooecial rows per branch over the duration of the Carboniferous Period. Zooecial apertures. The most significant information to emerge from the present investigation was that zooecial apertures could be assigned to three compact varieties as outlined below: 1. Fenestellid type — a simple, circular aperture with a narrow peristomal rim, the degree of exsertion of which was from weak to moderate depending upon the obverse branch profile. Mean apertural diameter was in the range of 0.08-0.14mm. The group contained no apparent internal apertural structures and included most but not all Carboniferous species previously referred to Fenestella by Australian authors. Further subdivision of this group seems quite possible when coupled with additional morphological features. 2. Polyporid type — a large, simple, circular or oval aperture with a very broad peristomal collar which may be either entire (Fig. 2, 5) or open proximally in a horse- shoe shape (Fig. 1, 4). Both conditions can be observed in the one specimen. External moulds of all members of the group reveal the presence of an operculum-like lid over the aperture. This cover bears either an elevated perforation or a spine-like projection, the exact nature of which is not evident in this form of preservation (Fig. 1, 7). The structure is usually placed in the centre of the lid but can be eccentric in some cases (Fig. 3, 8). Rare specimens have been observed to bear minute septa-like projections around the margin of the aperture as recorded in the type species Polypora dendrozdes M’Coy by Miller (1963). Mean apertural diameter is between 0.14-0.23mm. Species belonging to this group include most of the Australian Carboniferous forms previously referred to Polypora together with a small group of uncommon species which in all other aspects would have been reasonably placed in Fenestella. 3. Septatoporid type — a circular, strongly exserted to stalked aperture with a thin, high, calice-like peristome. Within the aperture there are eight vestibular septa radiating from a small central orifice. Mean apertural diameter is between 0.07- 0.13mm. Species of this group were previously placed in either Fenestella or Polypora depending upon the number of rows of zooecial apertures per branch. Apart from the apertures, other morphological aspects of this group leave little doubt of its separate status which has been elaborated elsewhere (Engel, 1975). With reference only to the fenestellid and polyporid types, it is significant that there is a separation of apertures into two size ranges with the larger forms bearing a low, wide, peristomal collar coupled with an opergulum-like covering over the orifice. It is also important that this separation does not conform with the conventional generic demarca.ion, based upon the number of rows of zooecial apertures per branch. The fact that some apertures bear traces of up to sixteen, weak, septal projections would support the contention (Engel, 1975) that there is an undoubtedly close relationship between the polyporids and the new genus Septatopora Engel. Zooecial chamber outline. The use of this feature as a diagnostic aspect for fenestrate genera is not without considerable risk of error. Wass (1968) has demonstrated that chamber outline is very sensitive to the exact level at which it has been recorded relative to the ‘basal plate’ of the specimen. Therefore, the use of random tangential thin sections for the identification of shape must remain a hazardous process, unless all shape changes have been recorded by serial sectioning. Morozova (1974) has expressed great confidence in the diagnostic value of chamber shape by using it with several other mesh characteristics to fragment the Proc. LINN. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL — 139 genus Fenestella (s.1.) into fourteen new genera. The present study does not entirely support the method adopted because of the difficulty in quantifying the actual shapes observed. When a chamber is studied as a three-dimensional internal mould, the distinction between triangular, trapezoidal and pentagonal shapes becomes a function of the branch width and the extent to which the chamber has been exposed down the sides of the branch. It is possible to recognize all three shapes on different branches within the one colony, given variable branch widths and different amounts of strip- ping of the outer calcareous layers. Thin sections cut at varying levels may record all of the above shapes, terminating with a bean-shaped outline when the section is deep enough to intersect the distal extension of the vestibule which leads to the obverse external aperture. Experience with Australian material would suggest that much more intensive serial sectioning of fenestrate species is necessary before zooecial chamber shape can be applied as a reliable diagnostic feature. There is no doubt, however, that the feature has a role to play in future classifications of fenestrate species. Nodes. In the species with three or more rows of apertures, described in this paper, nodes are frequently lacking. In the few cases where they are present, they occur randomly in any row being situated adjacent to the proximo-central rim of a particular aperture (Fig. 1, 8 & 9). This forms an interesting contrast to the contemporary Septatopora genus where the nodes are located on the distal rim of the apertures. Due to the width of the peristomal collar in the polyporids, it is not uncommon to see the node fused into the margin of the collar (Fig. 1, 8). The point to be noted is that there is a very close relationship of each node with a particular aperture and that there is not the development of a regular central row of nodes as recorded in the type species. Nodes in the dominantly two-rowed species described in this paper follow a regular, linear, nodal pattern when developed. Modes of preservation. One of the central difficulties experienced in working with fenestrate species concerns variation in the modes of preservation of specimens. Eastern Australian Carboniferous material is almost entirely preserved as external and internal moulds, with a consequent lack of primary skeletal material. Whilst this effectively prevents the preparation of thin sections, it undoubtedly provides excellent preservation of all external characteristics coupled with useful internal reconstructions of the original zooecial chambers. The few silicified specimens extracted from local limestones have a worn, open skeletal appearance which compares very badly with the surface details available from moulds. This is possibly due to incomplete replacement or post-depositional abrasion, for the criticism is not true of this style of preservation in other regions. A major problem arises in comparative studies because of different modes of pre- servation. Many Russian species are illustrated only with figures of thin sections which have been made at unspecified levels within the zoarium. It is impossible to compare these with the fine surface sculpture available from moulds. Comparative work is therefore reduced to the level of mesh comparisons which are, in themselves, insufficient for positive diagnosis in many cases. Undoubtedly these constraints of pre- servation have led to much duplication in the published literature. TAXONOMY Biologically, it would seem that the morphological details and dimensions of individual zooecia may have more relevance than the number of them that can be packed into an ever expanding branch width. This opinion needs substantiation from Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 140 FENESTRATE BRYOZOANS other workers since many older publications are deficient in this type of data, making literature comparisons very inconclusive. Pending wider consideration of the classification value of individual zooecial details as proposed in this paper, a relatively conservative taxonomic approach has been adopted. Instead of grouping all the species within one genus, two new genera are proposed namely Australopolypora gen.nov. and Australofenestella gen.nov. In the former, all multi-rowed, non-carinate species are grouped together into an evolving stream which permits reasonable comparison with the many existing species of Polypora M’Coy. Similarly, the dominantly two-rowed, carinate species are placed together in the second genus facilitating a ready comparison with species of Fenestella Lonsdale. Thus instead of being incompatible with the classical dichotomy based on rows of apertures, the new genera build on that basis and extend the definitions to include the apertural characteristics. This compromise reduced the strength of the argument that like apertures should be grouped together but it is the only realistic course to adopt in the confused field of fenestrate bryozoan taxonomy. It must be stressed however that the new genera exhibit a much closer relationship than the nomenclature would suggest. Reluctance to make major changes in the present generic status flows from the fact that the eastern Australian Carboniferous occurrence of fenestrate species is very restricted and is not necessarily representative of world-wide distribution. The present review of this fauna has revealed, after contractions by synonymy, a current total of fifteen species of Fenestella, seven species of Australofenestella, six species of Australo- polypora and nine species of the new genus Septatopora (Engel, 1975) together with less common examples of Archimedes, Hemitrypa and Semicoscinium. On the basis of such a small sample it is unwise to propose major taxonomic shifts. However, the mor- phological trends evident in this pilot study of a relatively uncomplicated fauna could possibly have wider implications for the considerably larger and more diverse faunas in other parts of the world. | Abbreviations. All abbreviations used in the statistical treatment of fenestrate mesh are of standard form as cited in Engel (1975, p.577). Repositorzes. Catalogued specimens have their number prefixed by the letter ‘F’, preceded by the following Museum coding: QU = Queensland University; QGS = Queensland Geological Survey; NEU = University of New _ England; NU = University of Newcastle; SU = University of Sydney. Fig. 1. (All except 6 prepared from latex casts.) 1-3. Australopolypora palenensts (Crockford). 1, 2, obverse surface showing wide, distally-inclined apertures with distinct, low, peristomal collars of similar orientation, QUF62187, locality Mt. Barney, x20, x30 respectively. 3, obverse surface showing similarly arranged apertures and peristomes, QUF32248, locality Ridgelands (188848), x20. 4-5. Australopolypora rawdonensis sp.nov. 4,5, obverse view of paratype showing horseshoe-shaped peristomes arranged in a fan-like form across the branch, NUF2345, locality NUL258, x15, x10 respectively. 6-9. Australopolypora altinodosa (Campbell). 6, reverse view showing shape and arrangement of internal moulds of three rows of zooecial cells, NEUF4708A, locality NUL9, x20. 7-9, obverse views of holotype illustrating: two-three zooecial row development; large open apertures with a boss-like projection on operculum; pseudo-carinal ornament in two-rowed branches; and large, broken, ribbed, surface spines, NEUF4708A, locality NULY, x20, x20, x10 respectively. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 141 B.A. ENGEL Tas ‘ ns Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 FENESTRATE BRYOZOANS 142 Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 143 Fossil localitzes. Where appropriate, localities are given by their number from the University of Newcastle Locality Index (NUL). Localities not in that index are given in full detail in the text. NUL9 — 3km east of Booral, N.S.W. (Campbell, 1961) ; NUL39 — Cameron's Bridge, Rouchel Brook, N.S.W. (Crockford, 1947) ; NUL258 — Barrington, N.S.W. (Cvancara, 1958) ; NUL454 — Isaacs Formation, Booral, N.S.W. (Campbell, 1961); NUL472 — Ridgelands 1 mile Military Map (Grid Reference 194827), Queensland (Fleming, 1969); NUL514 — Rawdon Vale, N.S.W. (Cobark 1:31680 Grid Reference 739454) ; NUL529 — Bonnington Forma- tion, Trevallyn, N.S.W. (Roberts, 1965). Photographic methods. Most specimens are preserved as internal or external moulds from which blackened latex casts were prepared for photographic purposes. These casts were then whitened with a layer of ammonium chloride before being photo- graphed with a camera attached to a stereobinocular microscope. SYSTEMATIC DESCRIPTIONS Order CRYPTOSTOMATA Shrubsole & Vine 1882 Family FENESTELLIDAE King 1850 Genus AUSTRALOPOLYPORA gen.nov. Type species. A. palenensis (Crockford) (= Polypora palenensis Crockford, 1949). Diagnosis. Unifoliate or funnel-shaped fenestrate expansions comprising a uniform mesh of radiating, straight or gently sinuous, usually non-carinate branches joined by regular, non-poriferous, transverse dissepiments; zooecial apertures in three or more rows on obverse branch surface with increase and reduction in number of rows at branch bifurcations; apertures with low, broad, entire or horseshoe-shaped peristome, scarcely raised above the branch surface; apertures with an operculum which bears a raised boss; nodes, if developed, situated on the proximo-central rim of an aperture; reverse surface may bear spiny projections; both obverse and reverse branches may be smooth, granular or longitudinally striate. Geological range. Late Visean — Permian. Remarks. The distinctive apertural form and nodal development of this genus set it apart from the polyphyletic genus Polypora M’Coy. Fig. 2. (All except 9 & 10 prepared from latex casts. ) 1-2. Australopolypora keppelensis parvula subsp.nov. 1, 2, obverse surface of holotype showing large, circular apertures with strong peristomal collars. Note nodal development and regular mesh, QUF32296, locality Ridgelands (194827) , x20, x40 respectively. 3-5. Australopolypora scalpta (Campbell). 3-5, obverse surface of holotype showing close packing of the large, circular apertures. Peristomal collars are depressed below branch level, NEUF4720B, locality NUL9, x3, x10, x30 respectively. 6-13. Australopolypora neerkolenstis (Crockford). 6, obverse surface of holotype, QUF25005, locality Mt. Barney, x10. 7, 11, 13 obverse surface illustrating low, peristomal collars surrounding wide apertures and sinuate longitudinal branch ornament. Note extended development of two zooecial rows after branch bifur- cation, QUF10893, locality Ridgelands (265791), x10, x10, x20 respectively. 8, obverse surface of specimen QUF10892, x10, locality. Ridgelands (265791). 9, 10, obverse view of eroded specimen showing erect vestibules leading up to the external apertures; and reverse view of zooecial cells showing irregularly penta- gonal to elongate rhomboidal form, QUF43162, locality Yarrol (QUL2654), x10, x15 respectively. 12, obverse view of specimen which has been flattened during preservation, NEUF5667, locality NUL390, x10. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 144 FENESTRATE BRYOZOANS Australopolypora rawdonensis sp.nov. Fig. 1, 4-5 Diagnosis. Coarse torm with wide branches and narrow dissepiments forming a rect- angular mesh; zooecia in three or four rows per branch with five to nine apertures per fenestrule; apertures oval, with horseshoe-shaped peristomes arranged in a fan- shaped arrangement across each branch. Description. Zoartum: Fan-shaped, being composed of radiating branches of unknown orientation; maximum radius 35mm. Obverse surface: (a) Branches. Very wide (m.BW 0.56mm), straight or broadly curved; branch cross section circular to slightly oval, commonly deflated; ornament of longitudinal, sinuous, pustulose ridges which meander around and between zooecial apertures. (b) Dzsseb¢ments. Narrow to medium width (m.DW 0.16mm) with only slight expansion at branch junctions; level with or below branches; ornament or ridges continuous on to branches. (c) Fenestrules. Sub-rectangular to rectangular; coarse, irregular, fan-shaped mesh; fenestrule openings equal to or greater than branch width resulting in an open meshed appearance; long, wide fenestrules (m.FL 2.71mm; m.FW 1.24mm). (d) Carina. Absent. (e) Nodes. Absent; a few specimens have rare, large, distant, irregularly spaced spines of presumed attachment significance. (f) Zooecial apertures. Medium size, oval (m.max.ZD 0.17mm); surrounded by a strongly pustulose, moderately elevated, horseshoe-shaped peristome, open on the proximal margin; aperture closed by a poorly-preserved, centrally-perforated plate. Oval-shaped apertures in central rows with long axis parallel to branch length; marginal rows with long axis at 45° to branch length, with peristome gap facing centre of branch, or drawn out into a proximally-directed club-shape. Apertures alternate in adjoining rows with strong marginal indentation of fenestrules but without dissepimental stabilization; zooecia in three or four rows per branch with increase to five pre-bifurcation and decrease to three post-bifurcation; apertures moderately spaced (m.Z-Z 0.37mm) with from 5 to 9 zooecia per fenestrule (m.Z/F 7.3). Reverse surface. (a) Form. Round branches joined by level or depressed, narrower dissepiments; reverse normally deflated; ornament of fine, longitudinal ridges com- parable with those of the obverse surface; several strong spines developed at base of zoarium. (b) Zooeczal bases. Oval bases arranged in non-overlapping rows. Materzal. Holotype NUF2343a/b (NUL258); Paratypes NUF2344, 2345, 2347, 2349a/b (NUL258); NUF2353 (NUL514); Others NUF2346, 2348 (NUL258) ; NUF2350, 2351, 2352, 2354, 2355 (NUL514). Remarks. No other Carboniferous species of Australopolypora has been found with a comparable apertural arrangement. Pustulose, horseshoe-shaped peristomes of similar form can be found at a lower stratigraphic level in such species as Fenestella allynensis Roberts (1965). The specific name is derived from the locality of Rawdon Vale, N.S.W. Stratigraphy. Of major significance is the fact that A. rawdonensis is the first develop- ment of a three-four rowed species in the Australian Carboniferous sequence. It is found to be of common occurrence in the Rhzpedomella fortimuscula zone at various localities in the Stroud-Gloucester Syncline, N.S.W. This stratigraphic distribution would indicate that Australopolypora first appeared in the Australian record in the mid-late Visean in marked contrast to the much earlier Northern Hemisphere distribution of Polypora M’Coy. Australopolypora palenensis (Crockford, 1949) Fig. 1, 1-3 1949 Polypora palenensis Crockford, pp.427-8, text fig. 12. 1972 “Polypora”’ palenensts Crockford; Fleming. pp.7-8, pl.3, figs. 1-3. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 145 Revised diagnosis. Coarse form with very wide branches and large sub-oval to rect- angular fenestrule openings; zooecia in three to four rows per branch, five to nine apertures per fenestrule; apertures circular, distant, inclined distally with proximal rim level with branch and distal rim depressed below branch level. Rewsed description. Zoartum: Fan-shaped, composed of radiating branches of unknown orientation; maximum radius 40mm. Obverse surface: (a) Branches. Very wide (m.BW 0.60mm) straight or broadly curved in an irregular pattern due to frequent bifurcation; cross-section circular to oval, but commonly deflated into a broad strap-like form; zooecial rows separated by faint longitudinal meandering ridges; interapertural spaces and sides of branch with similar sculpture. (b) Dissepiments. Slender to wide (m.DW 0.23mm); narrow forms have only slight expansion at branch junction; wide forms expand continuously from centre to branch junction resulting in variably-shaped fenestrules. (c) Fenestrules. Sub-oval to rect- angular; coarse, irregular, fan-shaped mesh; fenestrule openings large, resulting in an open meshed appearance; long to very long, wide fenestrules (m.FL 3.1mm; m.FW 1.37mm). (d) Carzna. Absent. (e) Nodes. Absent. (f) Zooeczal apertures. Medium size, circular (m. ZD 0.14mm), but frequently deformed during preserva- tion; peristome entire, uniform, low, but well developed, and inclined distally; apertures closed by a poorly-preserved, perforated plate. Zooecial chambers elongated proximally, with a distally-directed vestibule meeting the branch at a low angle, resulting in the apertural face also being distally inclined to the branch surface; this is achieved by having the proximal rim level with, or above the branch surface and the distal rim depressed into the branch in a suitable cavity; resultant longitudinal obverse branch profile is a sawtooth pattern along each row of apertures. Fenestrule indentation slight or lacking; apertures not stabilized with respect to dissepiments ; zooecia in three to four rows per branch with increase to five or six pre-bifurcation, and decrease to two or three post-bifurcation; apertures distantly spaced (m.Z-Z 0.48mm) with from five to nine zooecia per fenestrule (m.Z/F 6.4). Reverse surface: (a) Form. Rounded branches joined by narrower dissepiments generally situated below branch level; reverse commonly deflated; ornament of fine longitudinal striations normally obscured by a thin smooth overgrowth. (b) Zooeczal bases. Oval with little or no overlap between rows. Material. Holotype QUF25008, Neerkol Series, Por. 127V/202, Par. Palen, Mt. Barney, Queensland; Paratype QUF25009 (type area); Others QUF62187, QGSF10931la/b (type area) ; QGSF10909 (Neerkol Fm., Malchi Creek, Ridgelands 1 mile map ref. 265791); QUF32248 (Neerkol Fm., Ridgelands 1 mile map ref. 188848) . Remarks. Fleming (1972) amplified the brief description of Crockford (1949), and gave the first photographic illustration of the species. However, he omitted to emphasize the apertural inclination and its resultant obverse branch profile which, apart from the very coarse mesh, is the most diagnostic feature of the species. The low angle of intersection of the straight vestibule and branch surface has resulted in a dis- tinctive form which has not been observed in any of the other described species of Australopolypora. Small apertural septa noted by Fleming (1972) have not been observed in the material available to the writer. Some external moulds display a ring of very strong pustules around the peristome which are possibly equivalent to the tubules surrounding the apertures of P. dendrozdes M Coy. Stratigraphy. The majority of specimens of A. palenensis (Crockford) come from the Mt. Barney region or from Malchi Creek, near Rockhampton, Queensland. Lack of Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 146 FENESTRATE BRYOZOANS recognition of the species in other areas is probably a product of the brief description and inadequate type material upon which the species was based. Though not common in New South Wales, it has been found in the present study at a number of localities where it was not listed by previous authors. In all cases it occurs with a Levipustula levis fauna, but because of an inability to establish the time equivalence of many of the occurrences of this fauna, it is not possible to assign 4. palenensis a restricted range as Fleming (1972) has done for its occurrence in Queensland. Australopolypora altinodosa (Campbell, 1961) Fig. 1, 6-9 1961 Fenestella(?) altnodosa Campbell, pp.458-9, p1.57, figs. 2a,b. Revised diagnosis. Medium to coarse fenestrate with wide branches arranged in a sub- rectangular mesh; zooecia in two to three rows per branch, with three to four aper- tures per fenestrule; apertures large, distant with strong circular peristome; nodes very large and distantly spaced; branch ornament of strong ribbing. Revised description. Zoartum: Small, fan-shaped fragment of unknown orientation; maximum radius 20mm. Obverse surface: (a) Branches. Straight, wide (m.BW 0.46mm), commonly deflated; obverse branch profile centrally elevated but without median carina; ornament of strong, longitudinal ribbing, about six in number with central rib becoming carina-like on two-rowed branches; some ribs continuous and sinuate, others only developed interaperturally; ribs on lateral slope of branch bear fine pustules; rib number rises to twelve pre-bifurcation. (b) Dzssepements. Medium width (m.DW 0.17mm); centrally straight with moderate expansion at branch junction; situated level with branches; ornament of strong ribs continuous on to branches. (c) Fenestrules. Elongate sub-rectangular to sub-oval; medium length and width (m.FL 1.75mm, m.FW 0.93mm). (d) Carzna. Absent; role assumed by central prominent, sinuate, surface rib; early development of three rows of zooecia results in two such ribs over considerable distances. (e) Nodes. Very large, circular, oval-based with a diameter of 0.16-0.24mm; nodes very tall (20.4mm), generally sloping distally with neither obvious termination nor superstructure evident; ornament of longitudinal ribbing; distantly spaced (m.N-N 0.90mm), irregularly placed along a central row; nodes on three-rowed branches are situated between the apertures of the central row but seldom on the centre line of the branch. (f) Zooeczal apertures. Circular to slightly oval, large (m.ZD 0.20mm), surrounded by a prominent, moderately-exserted peristome; each aperture closed by an axially-perforated, domed, diaphragm-like plate; apertures alternate in lateral and median rows and are not stabilized with respect to the dissepiments; marginal rows with little or no fenestrule indentation; zooecia in two rows per branch with three rows developing up to 5mm prior to bifurcation; apertures distantly spaced in each row (m.Z-Z 0.49mm) with from three to four zooecia per fenestrule (m.Z/F 3.6). Reverse surface: (a) Form. Branches with broadly semi-circular profile tapering slightly from obverse; dissepiments level with or slightly below branches; ornament of fine, straight, longitudinal ribbing. (b) Zooeczal bases: Elongate, irregularly penta- gonal in marginal rows with the central row being rhomboidal in shape. Materzal. Holotype NEUF4708A, Booral, N.S.W. (NUL9). Remarks. Campbell (1961) noted several unusual features about this species which caused him to have doubts about its correct generic category. His final choice of doubtfully grouping it with Fenestella Lonsdale is at variance with other mor- phological aspects, here considered to place it more appropriately with Australopolypora. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 147 Unusual features noted by Campbell (1961) include: very high nodes placed randomly on any part of the obverse brarich surface, an absence of any clearly defined median carina, and the very early development of a third row of apertures up to 5 mm prior to each branch bifurcation. In addition to these aspects, the apertural form of the species is considered to align it closely with other Australian Carboniferous species of Australopolypora. It shares with them a very large circular aperture which is surrounded by a broad peristomal collar of distinctive appearance. It also exhibits the flat, centrally per- forated plate which is presumed to have closed the aperture. As discussed earlier, these apertural features are considered to be of greater diagnostic value than the number of rows of apertures which does not clearly group the species with either of the available generic categories. No comparable species has been described in available literature. Stratigraphy. The single known specimen was described by Campbell (1961) from just below the main Levpustula bed at Booral, N.S.W., a locality considered to be of early Westphalian age. No other comparable material has been recovered from the type locality or elsewhere. Australopolypora neerkolensis (Crockford, 1949) Fig. 2, 6-13 1949 P.neerkolensts Crockford, pp.426-7, text-fig. 10. 1962 P.neerkolensis Crockford, Campbell, pp.46-7, pl.13, figs. la-d. 1964 P.neerkolensts Crockford, Maxwell, p.58, [non pl. 13, figs. 8-9]. 1964 P. neerkolenszs Crockford, Hill & Woods, p.c.8, pl.C4, figs. 3-4. 1972 P.neerkolensts Crockford, Fleming, pp.5-6, pl.3, figs. 4-7. Revised diagnosis. Medium-sized form with very wide branches and wide dissepiments set in a variable mesh of oval to sub-rectangular fenestrules; zooecia in three rows per branch, with three to five apertures per fenestrule; apertures large, circular, with centrally-perforated domed plate; carina lacking; nodes irregularly developed. Rewsed description. Zoarium: Sub-parallel branches set in a narrowly-radiating fan- shaped zoarium of unknown orientation; maximum radius 50mm. Obverse surface: (a) Branches. Very wide (mBW 0.64mm) ; branch cross-section oval but commonly flattened by deflation, ornament of fine pustules with variable development of longitudinal ridges between apertures; ornament strong on branch sides; two-rowed branches can have a raised central rib of carina-like form. (b) Dzssepzments. Medium to broad (m DW 0.27mm) ; outline highly variable with some expanding continuously from their centre to the branch junction, whilst others are narrow and straight without expansion; most dissepiments are inclined rather than vertical in position; situated level with or just below branches; ornament of strong ribbing continuous on to the branch surface. (c) Fenestrules. Oval to sub-rectangular; medium-sized mesh of variable appearance; some specimens have fenestrule openings equal to or narrower than branches resulting in a closed mesh appearance whilst others have thin extended dissepiments which produce an open-meshed form; medium length, medium to wide fenestrules (m FL 1.74mm; m FW 1.10mm). (d) Carina. Absent in all but short post-bifurcation segments where a central prominent rib can develop a carina-like appearance between the two rows of apertures which are present in that region. (e) Nodes. Irregularly developed nodes occur on some specimens, being placed adjacent to, or forming part of, the apertural peristome on its proximal side in any apertural row. (f) Zooecial apertures. Large (m ZD 0.20mm), circular to oval, being surrounded by a prominent, entire peristome; apertures crossed by an arched, centrally perforated plate which in external moulds bears a raised central boss which Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 148 FENESTRATE BRYOZOANS can rise above the level of the peristome; usually three rows per branch with increase to four or five pre-bifurcation and decrease to two or three rows post-bifurcation ; apertures distantly spaced in each row (m Z-Z 0.44mm) with from three to five apertures per fenestrule (m Z/F 4.3) ; apertures in adjoining rows very closely packed with less than a zooecial diameter separating them diagonally; apertures may indent fenestrule margin according to the degree of branch deflation, and are not stabilized with respect to dissepiments. Reverse surface: (a) Form. Narrowly-rounded branches joined by level or slightly depressed dissepiments; thin outer surface smooth beneath which are developed numerous fine, longitudinal striations. (b) Zooeczal bases. Elongate oval to elongate rhomboidal ; irregularly pentagonal in two-rowed segments. Material. Holotype QUF25005, Por.127V/202 Par. Palen, Mt. Barney; (Paratype — Specimen QUF24999 listed by Crockford (1949) as a paratype is actually a specimen of Septatopora pustulosa). Other specimens mentioned by Crockford are missing; Others QGSF10929-10930, 10934 (Type locality) ; QGSF10892-10894, Malchi Creek, Stanwell; QUF32189 Neerkol Creek, Stanwell; QUF32290, NUF2367-2368 (NUL472) ; QUF43162, 43199 Yarrol (Maxwell, 1964); NEUF5656-5670 Oaky Creek (Campbell, 1962). Remarks. Crockford (1949) based this species on a poorly-preserved holotype which has resulted in considerable confusion with later determinations. Better-preserved material described by Campbell (1962) and Fleming (1972) has resulted in a clearer understanding of the species. In the present study, considerable doubts have been entertained with respect to the conspecific nature of all the material placed by Fleming (1972) in this species. Features which exhibit most variation include: (a) Mesh form. Most specimens have broad, curved dissepiments associated with a closed-mesh appearance. A few specimens (QGSF10892-4) have thin, extended dissepiments with a resultant open mesh. This variation could be due to meshwork changes within a complete zoarium, but larger specimens from other regions do not support such a proposal. (b) Zooeczal apertures. Well preserved branches have a round profile with moderately-exserted apertures which indent the fenestrule margin, thus having a somewhat similar appear- ance to that of Septatopora pustulosa (Crockford) with which it was confused by both Crockford (1949) and Maxwell (1964). Many other specimens have strongly-deflated, strap-like branches in which the apertures are also flattened. The difference in appearance between these two states of preservation requires a very detailed inspection for transitional stages between the two extremes. (c) Apertural diameter. Open- meshed specimens (QGSF10892-4) have a consistently larger zooecial diameter than that of most other specimens. The combination of this feature plus the mesh form may be significant, but, as it is possible to observe transitional stages, some uncertainty persists as to the correct specific designation. The holotype is most certainly too badly preserved to enable a positive diagnosis of the morphological limits of the species. Stratigraphy. A. neerkolensis occurs in the Levipustula levis zone at various localities in New South Wales and Queensland. Queensland specimens have been recovered from various levels in the Neerkol Formation (Stanwell), from the Poperima Formation (Yarrol), and from the type locality at Mt. Barney. New South Wales specimens have been recorded from the Kullatine ‘Series’ at Oaky Creek (Campbell, 1962) and from several localities of the Levipustula levis assemblage in the Gloucester-Bulahdelah region. Fleming (1972) suggests that A. neerkolenszs is restricted to the early portion of the Leuipustula levis zone in the Neerkol Formation and its equivalents. Re- examination of specimens placed by Fleming (1960, 1969) in Polypora cf.woods: from Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 149 the Aurtculispina levis zone, together with extra material collected from that zone (QUF32290, NUF2367, NUF2368) leave little doubt that they belong to 4. neerkolensis. In these circumstances it would appear that this species is a long ranging one extending through the whole of the Neerkol Formation (2100m). Australopolypora scalpta (Campbell, 1961) Fig. 2, 3-5 1961 P.scalpta Campbell, pp.461-2, pl.60, fig. 11. Revised diagnosis. Medium-sized form of closely-meshed appearance; composed of very wide branches and very small, oval fenestrules; carina and nodes absent; apertures large, oval, medium-spaced being located in branch depressions from which a thin, entire peristome rises almost to branch level; apertures usually in four rows per branch. Revised description. Zoartum: Gently-radiating branches of unknown orientation; maximum radius 12mm. Obverse surface: (a) Branches. Very wide (m.BW 0.86mm), generally flattened in cross-section; ornament of sinuous, longitudinal ridges between apertural rows; surface pustulose. (b) Dzssepiments. Very wide (m.DW 0.74mm); strongly expanded at branch junction; level with branches; ornament of some surface ribbing, more evident on sides of both branches and dissepiments. (c) Fenestrules. Oval; very closely-meshed form dominated by wide branches and small fenestrules, resulting in a very closed mesh appearance; medium length, medium to wide fenestrules (m.FL 1.56mm; m.FW 1.14mm); actual fenestrule opening is approximately 0.8mm long by 0.3mm wide. (d) Carzna. Absent; three to five linear rows of apertures are separated by prominent, sinuous ridges. (e) Nodes. Absent. (f) Zooeczal apertures. Large, slightly oval (long diameter m.ZD 0.23mm) ; surrounded by a narrow, raised, entire peristome which does not quite rise to the level of the longitudinal, interzooecial, sinuous ridges, due to the moderate depression of the apertures below branch level; apertures in four rows per branch with increase to five pre-bifurcation and decrease to three post-bifurcation; apertures moderately to distantly spaced (m.Z-Z 0.41mm) with from three to four zooecia per fenestrule (m.Z/F 3.8) ; apertures in adjoining rows closely crowded, being separated diagonally by a distance of about half the zooecial diameter ; fenestrules not indented by apertures which are also not stabilized with respect to the dissepiments. Reverse surface: (a) Form. Flattened to weakly-rounded branches joined by broad, level dissepiments ; branches bear weak, finely pustulose striations usually obscured by secondary thickening. (b) Zooeczal bases. Broadly flattened, elongate-hexagonal or oval in outline. Material. Holotype NEUF4720A/B, Booral, N.S.W. (NUL9). Remarks. This highly distinctive species is readily recognizable by its depressed aper- tures and peristomes which are situated below the prominent, longitudinal, sinuous ridges separating the zooecial rows. No other comparable form has been observed in the literature. The holotype remains the only known specimen despite intensive collecting at the type locality. Stratigraphy. The type locality was recorded by Campbell (1961) as being above the main Levipustula levis bed at Booral, New South Wales (NUL9). Australopolypora keppelensis (Crockford, 1946) 1946 P. mznuta Crockford, p.133, text fig. 9. [non] 1932 P. minuta Deiss, p.28. [| vde Crockford, 1962]. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 150 FENESTRATE BRYOZOANS 1962 P. keppelenszs Crockford, p.840. 1968 P. keppelenszs Crockford, Wass, p.47, pl.12, fig.2. Diagnosis. Medium to fine form; zooecia in three rows with three zooecia opposite each fenestrule; surface ornamented by discontinuous ridges and grooves between the apertures, and by a few small nodes. Holotype. QUF7974a, Lakes Creek Beds (Trachypora horizon behind quarry) , Rock- hampton, Queensland. (Artinskian) . Geological age. Late Carboniferous — late Permian. Australopolypora keppelensis parvula subsp.nov. Fig. 2, 1-2 Diagnosts. Medium to fine species; regular mesh with wide branches and dissepiments forming uniform oval fenestrules; carina absent; nodes irregularly disposed adjacent to some apertures; apertures large, closely spaced in three rows per branch; peristome wide being either entire or proximally open; apertures closed by a flat, centrally perforated plate. Description. Zoartum: Moderately-radiating, sub-parallel branches in flat expansion of unknown orientation; maximum radius 22mm. Obverse surface: (a) Branches. Medium to wide (m.BW 0.41mm) with circular cross-section becoming oval at bifur- cation; ornament of sinuous ribbing between and around the apertures. (b) Dissepiments. Wide (m.BW 0.23mm), expanded from centre in a semicircular curve to the branch junction; level with branches; ornament of strong ribbing continuous on to branches. (c) Fenestrules. Oval to sub-oval; mesh fine to medium-sized and very regular; fenestrule openings and branch width of similar dimensions producing a very uniform mesh appearance; fenestrules short and of medium width (m.FL 0.84mm; m.FW 0.64mm). (d) Carina. Absent. (e) Nodes. Numerous, round (diam.ca. 0.1mm), bluntly-pointed, irregularly-developed nodes associated with apertures in all zooecial rows, but tending towards a near central arrangement; absent over large areas, but when present usually placed adjacent to the proximal rim of an aperture. (f) Zooeczal apertures. Medium to large (m.ZD 0.15mm), circular to oval in outline; peristome prominent, slightly raised and faintly pustulose; peristome can be entire, but more frequently has a horseshoe-shaped appearance, with the proximal margin being smoothed over in a lip-like form; each aperture covered by a centrally-per- forated plate; apertures in three rows per branch with increase to four pre- bifurcation, and decrease to two post-bifurcation; fenestrule indentation slight, and apertures not stabilized with respect to the dissepiments; apertures directed perpen- dicular to the curvature of the branch with some marginal elevation of the peristomes in the lateral rows; branch surface depressed between apertures which are closely spaced (m.Z-Z 0.26mm) with from two to four zooecia per fenestrule (m.Z/F 3.2) ; apertures in adjoining rows very close being diagonally separated by about half a zooecial diameter. : Fig. 3. (All except 6 prepared from latex casts. ) 1-2. Australofenestella brookeri sp.nov. 1, 2, obverse surface of holotype showing large apertural form in a two-rowed species. Some apertures exhibit a central axial boss, NUF2541, x15, x30 respectively. 3. Australofenestella stroudensis minuta subsp.nov. 3, obverse surface of holotype showing large apertural form and a low, central, nodose carina, NUF2396, locality NUL258, x20. 4-9. Australofenestella stroudensis stroudensis (Campbell). 4, obverse surface of specimen previously referred to Fenestella anodosa Campbell, NEUF4701, locality NUL9, x20. 5, obverse surface of holotype of F. anodosa, NEUF4700C, locality NUL9, x20. 6, reverse surface of holotype of A. s. stroudenszs showing arrangement of zooecial cells, NEUF4704B, x15, locality NUL9. 7-9, obverse surface of holotype of 4. s. stroudensis illustrating the large apertural form and distinct development of a strong boss or spine on the operculum covering the apertures, NEUF4704B, x20, x30, x10 respectively. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 151 B.A. ENGEL Proc. LINN. Soc. N.S.W., 103 (3), (1978) 1979 FENESTRATE BRYOZOANS 152 Proc. LINN. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 153 Reverse surface: (a) Form. Rounded branches joined by medium-width dissepiments slightly below branch level; ornament of a smooth, thin, outer layer covering longitu- dinal striations. (b) Zooeczal bases. Irregularly pentagonal in lateral rows with a rhomboidal form in the central rows. Materzal. Holotype QUF32296 (NUL472); Paratypes QUF32297, QUF32295 (NUL472). Remarks. The specimens which form the basis of this new subspecies were collected by Fleming (1960) who gave an unpublished description of them under the open nomen- clature of Polypora cf. keppelensis Crockford. The same material was listed again in this form in Fleming (1969). Fleming considered his material to be identical with A. heppelensis except for their consistently shorter fenestrule length. An examination of measurements made upon the type specimens of A. &. keppelensts and the specimens of A. k. parvula, given in Table 1, reveals that there are significant differences in fenestrule length and width, dissepiment width and zooecial spacing with consequent discrepancies in the associated space counts. Apart from these mesh differences, both groups display considerable similarity in apertural form and arrangement with some apertures of both having a proximal break in the peristome with an associated lip-like structure. Based upon the present material from the late Carboniferous Neerkol Formation as well as that from the Artinskian Lakes Creek Beds (Crockford, 1946) and from younger Permian beds in the Bowen Basin (Wass, 1968), the following stratigraphic changes support the erection of the new subspecies: (a) there is a noticeable change in mesh dimensions from 4. k. parvula in the Neerkol Formation to 4. k. keppelenszs from the Lakes Creek Beds with only slight further change in the later Permian specimens. (b) With the mesh change, the fenestrule form also changes from the regular oval shape in 4. k. parvula to sub-oval to sub-rectangular in 4. k. keppelensis from the Lakes Creek Beds to a normally sub-rectangular form in the younger A. &. keppelensis from the Bowen Basin. (c) Nodes of irregular distribution are very common in A. k. parvula, becoming rarer and finally absent in the Permian representatives. Stratigraphy. All specimens of A. k. parvula were collected by Fleming (1960) from the locality NUL472 (QGSL1000) as geologically described by Fleming (1969). This location, east of the Ridgelands-Stanwell Road, occurs in the top 300m of the Neerkol Formation and has been assigned a late Carboniferous age. A. k. keppelensts Crockford (1946) was originally described from the Lakes Creek Beds, east of Rockhampton. Because of breaks in the sequence, the section Fig. 4. (All except 3 & 6 prepared from latex casts. ) 1-3. Australofenestella(2) keepitensts sp.nov. 1, 2, obverse surface of holotype illustrating the close, irregular nodal distribution and the occasional development of a third central row of apertures. Note nodal variation (whiter marks) in 2 with both linear and zig-zag form. 3, obverse fragment of holotype showing three-rowed zooecial arrangement, NEUF7466A, locality NEUL318, x6, x30, x10 respectively. 4-9. Australofenestella malchi (Crockford). 4, 5, obverse surface of holotype showing large apertures, peristomal collars and large surface hemispherical depressions, QUF24952, locality Malchi Creek, x20, x20 respectively. 6, reverse view of zooecial chambers, QUF10903, locality Malchi Creek, x20. 7, obverse surface of specimen previously referred to Fenestella cerva Campbell. Note carinal development at branch bifurca- tions, NEUF4705H, locality NUL454,. x20. 8, 9, obverse views of holotype of F. cerva illustrating profile change as a result of deformation. Note boss-like projections within the apertures on 9, NEUF4709A, locality NUL454, x20, x20 respectively. 10, 11. Australofenestella cioncta (Crockford). 10, obverse surface of deformed specimen showing strong boss-like structures within the peristome, NEUF4715E, locality NUL9, x10. 11, obverse surface of well preserved specimen, QGSF10897, showing strong nodeless carina separating two rows of large apertures, locality Ridgelands (265791), x10. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 154 FENESTRATE BRYOZOANS connecting these two faunas is not known. On other faunal evidence, the Lakes Creek Beds were assigned an Artinskian age by Crockford, a determination supported by Kirkegaard, Shaw & Murray (1970). Genus AUSTRALOFENESTELLA gen.nov. Type species. Australofenestella malcht (Crockford) (=Fenestrellina malchi Crockford, 1949, pp.422-423, fig. 4) Diagnosis. Unifoliate or funnel-shaped fenestrate expansions comprising a uniform mesh of radiating, straight or gently sinuous, weakly to strongly carinate branches joined by regular, non-poriferous dissepiments; large size zooecial apertures in two rows with variable increase in rows prior to bifurcation; apertures with low, broad, entire or horseshoe-shaped peristome raised only on branches with acutely triangular cross-section; apertures with an operculum which bears a raised boss; nodes vary from obsolete to blunt, widely-spaced cones arranged in a central row on the carina; obverse and reverse branch surfaces may be smooth, granular or striate. Geological range. Carboniferous- Permian. Remarks. The distinctive apertural form in this genus readily separates it from Fenestella Lonsdale. Features which separate it from Australopolypora include the number of apertural rows, the median carina and the different arrangement of the nodes. Australofenestella brookeri sp.nov. Fig. 3, 1-2 Diagnosis. Medium-sized form with medium-width branches forming a sub-oval to sub-rectangular, regular mesh; branches bear no distinct central carina but a centrai row of large, broadly cone-shaped, distantly-spaced, nodes; apertures occur in two rows with a third row appearing only in the fork at bifurcation; apertures large, close to medium-spaced with a low, wide, entire peristome; zooecial bases elongate tri- angular to irregularly pentagonal. Description. Zoartum. Gently expanding fragments of unknown orientation; maximum radius 30mm. Obverse surface: (a) Branches. Medium width (m.BW 0.33mm) near straight; branch cross-section broadly rounded without carina; no ornament observed. (b) Duzussepiments. Medium width (m.DW _ 0.17mm) ; dissepiments expand in a broad curve from their centre to the branch junction where they join just below branch level; ornament of moderate ribbing which continues on to branch sides. (c) Fenestrules. Sub-oval to sub-rectangular; medium-sized, moderately regular mesh (m.FL 1.49mm; m.FW 0.72mm). (d) Carina. Absent; some indistinct low ridging occurs between nodes. (e) Nodes. Wide-based, cone- shaped nodes which narrow rapidly to a fine point; distantly spaced (m.N-N 0.71mm) in a central row. (f) Zooeczal apertures. Medium to large, circular (m.ZD 0.15mm); surrounded by a low, entire, broad, moderately raised peristome; apertures arranged in two rows with a third appearing only in the fork at each bifurca- tion; apertures alternate in adjoining rows and are situated on the broad obverse slope of the branch where they are directed with slight lateral inclination towards the fenestrule; they are not stabilized with respect to the dissepiments and have moderate fenestrular indentation; apertures close to medium-spaced (m.Z-Z 0.29mm) with from 4 to 6 zooecia per fenestrule (m.Z/F 5.1). Reverse surface. (a) Form. Unknown. (b) Zooeczal bases. Elongate triangular to irregularly pentagonal. Material. Holotype NUF2541 (NUL39). Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 155 Remarks. Diagnostic aspects of this species inciude its apertural form and position, branch form and nodes. The need to document these distinctive, rare elements of the Australian Carboni- ferous fenestrate fauna justifies the generally undesirable practice of erecting a new species based upon a single specimen. : The specific name honours a resident of the Rouchel Brook district, Mr B. Brooker. A. brookeri shares with A. stroudensis minuta and A. stroudensts stroudenszs the development of large cone-shaped nodes, wide, circular apertures, no carina, and a similar broad branch profile. Further, in order of stratigraphic appearance, A. brookert has one aperture in the fork at branch bifurcation, A. s. mznuta has two apertures and A. s. stroudensis can have between five and ten extra apertures in a third row prior to a branch division. This morphology would suggest that these three taxa form a lineage throughout the Carboniferous sequence. Stratigraphy. The only known material has been discovered in the Waverley Forma- tion at Cameron’s Bridge, Rouchel Brook, where it is associated with a fauna of the Pustula gracilis subzone of the Schellwienella cf. burlingtonensts zone. Australofenestella stroudensis (Campbell, 1961) 1961 F. stroudensts Campbell, p.458, pl.56, figs. la-c. 1961 F. anodosa Campbell, p.457, pl.57, figs. 3a-c. Revised diagnosis. Fenestrate species with medium to coarse, regular mesh; carina low, poorly defined with moderate to distantly spaced nodes of variable form; two rows of apertures increase to three some distance before branch bifurcation; apertures large, with entire peristome; branch ornament of longitudinal ribbing; zooecial bases irregularly pentagonal. Geological range. Late Visean- Westphalian. Australofenestella stroudensis stroudensis (Campbell, 1961) Fig. 3, 4-9 Synonymy. As above. Revised diagnosis. Medium to coarse species with wide branches arranged in a regular mesh; carina low, ill-defined, with distant, poorly developed obsolete nodes; apertures in two rows per branch with the appearance of a third row, containing up to ten apertures, before bifurcation; apertures large, distant, with entire circular peristome; branch ornament of sinuous, pustulose ribbing; zooecial bases irregularly pentagonal. Description. Zoartum: Expanding zoarial fragments of unknown orientation; maximum radius 60mm. Obverse surface: (a) Branches. Straight, wide (m.BW 0.45mm), commonly deflated; obverse branch profile centrally elevated with a low, ill-defined carina; ornament of sinuous, longitudinal, pustulose ribbing between and surrounding apertures. (b) Dzssepiments. Medium width (m.DW 0.17mm) ; cen- trally straight with moderate expansion at branch junction; situated level with branches; ornament of coarse to fine ribbing which expands laterally on to branch sides. (c) Fenestrules. Sub-rectangular, medium to coarse, moderately regular mesh; fenestrules medium to long and wide (m.FL 1.91mm; m.FW 0.97mm). (d) Carina. Low, rounded to blunt keel of variable preservation due to frequency of branch deflation; it may appear as a high, bladed structure if branches are depressed on either side of this central plate or alternatively, if the whole branch is flattened, it may be barely apparent on the obverse surface; two carinae develop with the very early Proc. LINN. Soc. N.S.W., 103 (3), (1978) 1979 156 FENESTRATE BRYOZOANS appearance of a third row of zooecia before bifurcation; because of the very large apertures, the carina tends to be sinuate or resorbed by the apertures. (e) Nodes. Some regions on the type material display low, indistinct, rounded elevations which may be taken to represent an obsolete row of nodes. (N-N. ca. 0.60mm). (f) Zooeczal apertures. Large, circular, prominent (m.ZD 0.19mm); slightly raised apertures surrounded by an entire peristome which may rise on to or replace the low carina; apertures arranged in two rows with a third row, containing from two up to ten aper- tures (max. length 5mm), appearing prior to each branch bifurcation; apertures alternate in adjoining rows and are partially stabilized with respect to the dissepi- ments; apertures placed on obverse branch surface, being either erect or gently inclined towards the fenestrule, unless distorted by preservation; fenestrular margin very slightly indented, if at all; apertures medium to widely spaced (m.Z-Z 0.43mm) with from 4 to 5 zooecia per fenestrule (m.Z/F 4.5). (g) Additzonal features. Zooecial diameter at the peristome is maintained vertically for the length of the vestibule, giving the appearance of a very wide, open aperture; at the base of this deep, erect vestibule the chamber turns in a sharp right-angle bend in the proximal direction, where it expands into an elongate zooecial chamber. A few branches display spherical depressions on the branch surface between apertures which may possibly represent the site of ovicellular development. Reverse surface: (a) Form. Rounded branches joined by narrow, level dissepiments; bifurcations preceded by a very gradual spread in branch width associated with the obverse development of a third row of apertures; many branches exhibit deflation effects on the reverse; ornament of longitudinal striations on branches and dissepi- ments, partly obscured by overgrowth in proximal parts of the zoarium; some large anchoring spines occur near the base of a few zoaria. (b) Zooeczal bases. Irregularly pentagonal. Material. Holotype NEUF4704A/B (NUL9) ; Paratype NEUF4703 (NUL9) ; Others NEUF4700B/C, 4701 (NUL9). Remarks. Detailed comparison of A. stroudensss (Campbell) and A. anodosa (Campbell) has revealed that most apparent differences are the result of the highly distorted state in which the type materials have been preserved. Type specimens of A. stroudenszs have been subjected to vertical compression which has produced a general flattening of the obverse surface. Specimens of 4. anodosa have been deformed by lateral compression in addition to the vertical loading. Whilst general deflation of the branch has taken place, the carinal plate has not been crumpled so that it now appears as a high, usually inclined, bladed carina. On either side of this carina, the weaker apertural surface has collapsed into a channel located between the carina and the doubled-over side walls of the branch, which now appear as two lateral carinae. Dissepiments are generally undeformed, but are snapped off at the branch junction, and pushed up over the apertural surface or into the side of the branch. Restoration of the profile would result in a form indistinguish- able from that of A. stroudensis. Both species were originally described as being nodeless. However, intensive study has revealed traces of very poorly preserved, low nodes on a few branches, a conclusion already noted by Wass (1968, p. 83). In both cases it is apparent that the nodes were very close to obsolescence. The existence of stronger, but similar nodes upon the strati- graphically older A. stroudens?s mznuta would support this conclusion. The only mesh variation between A. anodosa and A. stroudensis occurs in the zooecial spacing which is slightly greater in A. anodosa. This difference is not considered to be significant. In strict order of appearance in the one paper, A. anodosa is the senior specific Proc. LINN. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 157 name, but because it is considered inappropriate to apply this name to a species which does in fact bear nodes, A. stroudensis has been selected as the more suitable, available taxon. Stratigraphy. All known specimens have been found associated with the Lew/pustula levis zone at Booral, N.S.W. (Campbell, 1961). Australofenestella stroudensis minuta subsp.nov. Fig. 3,3 Diagnosis. Medium-sized fenestrate, with narrow branches arranged in a regular, open, sub-rectangular mesh; branches bear a low, ill-defined carina with small, moderately spaced nodes; apertures in two rows, with a third row appearing one or two apertures before bifurcation; apertures large, closely spaced with thin, entire peristomes; branch ornament of moderately developed longitudinal ribbing; zooecial bases irregularly pentagonal. Description. Zoartum: Moderately expanding, laminar, zoarial fragment of unknown orientation; maximum radius 30mm. Obverse surface: (a) Branches. Straight, narrow (m.BW 0.29mm) ; branch cross-section rounded with a gentle slope on either side of an ill-defined carina; ornament of moderately developed ribbing or otherwise smooth. (b) Dessepiments. Narrow (m.DW 0.08mm) ; centrally straight with greatest expansion at branch junction; situated level with branches and ornamented with coarse ribbing which expands laterally on to branch sides. (c) Fenestrules. Sub- rectangular; regular, open mesh with fenestrules of medium length and width (m.FL 1.37mm; m.FW 0.72mm). (d) Carina. Low, rounded, ill-defined central carina, partly resorbed by apertural peristomes. (ce) Nodes. Central row of small, circular or oval-based nodes with moderate spacing (m.N-N 0.44mm). (f) Zooeczal apertures. Large, circular, prominent (m.ZD 0.17mm) slightly raised apertures surrounded by a thin, entire peristome which can replace portion of the carina; wide vestibular dia- meter is maintained until further expansion occurs into the zooecial chamber; apertures arranged in two rows per branch, with a third row appearing up to two apertures prior to branch bifurcation; apertures alternate in adjoining rows and are situated on the broad obverse slope of the branch, being directed upwards or with slight lateral inclination; apertures partly stabilized with respect to dissepiments and only with slight marginal indentation of the fenestrules; apertures closely spaced (m.Z-Z 0.30mm) with from 4 to 6 zooecia per fenestrule (m.Z/F 4.6). Reverse surface. (a) Form. Broadly rounded branches joined by narrow, near level dissepiments; ornament of fine, longitudinal ribbing. (b) Zooeczal bases. Irregularly pentagonal. Materzal. Holotype NUF2396 (NUL258) . Remarks. The sub-specific name alludes to the diminutive size of this specimen as compared with that of A. s. stroudensis. Principal differences at the subspecific level include: (a) A. s. mznuta has an overall smaller mesh, with significant differences in most dimensions. Whilst size alone is not of major importance, the lack of intermediate material makes it unwise to group these specimens into a single taxon; (b) A. s. mznuta has a regular, central row of small nodes not developed on A. s. stroudens?s, where the nodes are either poorly pre- served or vestigial in form; (c) A. s. mznuta has fewer pre-bifurcation apertures than A. s. stroudensis. Generally A. s. minuta is closely related to A. s. stroudensis but displays sufficient transitional differences between A. brookerd and A. s. stroudenses to justify its treatment as a separate subspecies. Differences between A. s. mznuta and A. brookerz are noted in the description of the latter species. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 158 FENESTRATE BRYOZOANS Stratigraphy. This specimen has been collected from the Barrington area (NUL258) where it is associated with a Rhzpzdomella fortemuscula fauna (Cvancara, 1958). Australofenestella trevallynensis sp.nov. Fig. 5, 4-8 Diagnosis. Medium-sized, irregular, crenulated fenestrate with narrow to medium width, straight or broadly curved, weakly carinate branches forming a rectangular to sub-rectangular mesh; nodes medium size, moderately to distantly spaced; apert--res medium to large size, not stabilized, directed obversely or slightly sideways; zooecia in two rows with a third row appearing up to two apertures before bifurcation; zooecial bases elongate triangular/trapezoidal trending to irregularly pentagonal on wide branches. Description. Zoarium: Expanding, crenulated fragments of unknown orientation; maximum radius 60mm. Obverse surface: (a) Branches. Narrow to medium width (m.BW 0.30mm), straight or slightly irregular; radiating proximally, sub-parallel distally; cross-section broadly tapered to rounded but frequently deformed ; ornament of strong, pustulose, longitudinal ribbing often obliterated by smooth overgrowth. (b) Dissepiments. Narrow to medium width (m.DW 0.15mm); centrally straight with moderate expansion at branch junction; some junctions inflated to house an enlarged zooecium; ornament of strong ribbing with central rib having a carinate form. (c) Fenestrules. Rectangular to sub-rectangular; irregular mesh with fenestrules of medium length and width (m.FL 1.57mm; m.FW 0.73mm). (d) Carina. Moderately to weakly developed, low carina connecting elongated nodal bases; generally about one third of branch width. (e) Nodes. Medium-sized, pointed, circular nodes with elongated bases; medium to distantly spaced (m. N-N 0.59mm) in a central row. (f) Zooeczal apertures. Medium to large (m. ZD 0.14mm) ; circular to slightly oval-shaped ; one specimen has a few enlarged zooecia (ca. 0.2mm) located at branch-dissepiment junctions; peristome circular, slightly raised on fenestrular margin and covered with an operculum-like plate which bears a central boss; _apertures in two unstabilized rows with a third row of one or two apertures prior to bi- furcation; apertures closely spaced (m. Z-Z 0.31mm) with from four to six apertures per fenestrule (m. Z/F 5.1). Reverse surface: (a) Form. Evenly rounded or slightly tapered branches joined by medium-width dissepiments; ornament of fine striations with weak pustules usually rendered smooth by overgrowth. (b) Zooeczal bases. Elongate triangular/trapezoidal trending to irregularly pentagonal on wider branches. Fig. 5. (All except 8 prepared from latex casts. ) 1-3. Australofenestella cincta (Crockford). 1, 2, obverse surface of neotype showing large mesh form and very high, nodeless carina; 2 illustrates the development of large hemispherical pits superimposed on some apertures, within which it is still possible to observe the boss-like projection, QGSF10898, locality Ridgelands (265791), x10, x25 respectively. 3, obverse view of one branch showing the strong boss-like projections in each aperture, NEUF4715E, locality NUL9, x20. 4-8. Australofenestella trevallynensis sp.nov. 4-7, obverse surface of holotype showing large apertural form, peristomal collars and an ill-defined carina with evenly spaced, distant nodes, NEUF6918, locality NUL529, all figs. x20. 8, reverse of zooecial cells showing triangular-trapezoidal form, NEUF6908, locality NUL529, x20. 9-11. Australofenestella macleayensis (Campbell) . 9,11, obverse surface of holotype showing large apertures separated by a strong carina upon which the nodes are very difficult to observe, NEUF5738, locality NUL390, x10, x5 respectively. 10, obverse surface of holotype of A. crockfordae (Campbell) showing the typical crushed appearance of the material placed in that species, NEUF4699A, locality NUL9, x10. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 159 B.A. ENGEL Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 160 FENESTRATE BRYOZOANS Materzal. Holotype NEUF6918a/b (NUL529); Paratype NEUF6908/6909 (NUL529); Others NUF2429a/b, 2433 (NUL258); NEUF6918, (?)6919 (NUL529). Remarks. Features including carinal form, branch profile and apertural form display considerable variation in this species, largely as a result of structural deformation during preservation. Specimens from NUL258 (Barrington) tend to have little carinae, more exserted apertures and occasional enlarged zooecia whereas those from the type locality are more carinate and less exserted. The specimens from NUL529 (Trevallyn) were formerly grouped by Roberts (1965) in F. allynenst’s Roberts. They can be readily separated from the latter species which has large, ear-like apertural hoods, rounded carinae, much closer nodes and irregularly pentagonal zooecial bases. A. trevallynensis is quite similar to F. gresfordensis Roberts (1963) which is dis- tinguished by its possession of much closer nodes; distinct, high-bladed carinae; weakly-hooded, proximally inclined apertures; and very distinctive, flanged, zooecial bases. Stratigraphy. This species has been identified from the Trevallyn locality of Roberts (1965) in a formation containing an Orthotetes australis fauna. It has also been found in the Rhzpidomella fortiémuscula zone at Barrington (Cvancara, 1958), but not as yet in the intervening Delepinea aspinosa zone. Australofenestella malchi (Crockford, 1949) Fig. 4, 4-9 1949 Fenestrellina malchi Crockford, pp.422-423, fig. 4. 1961 Fenestella malchz (Crockford) Campbell, p.460. 1961 F. cerva Campbell, pp.455-456, pl.59, figs. la-c. 1964 F. malchz (Crockford) Maxwell, p.38, pl.12, fig. 13. 1972 F. malchi (Crockford) Fleming, p.4, pl.2, figs. 1-3. Revised diagnosis. Regular fenestrate with medium-width, straight branches forming a uniform sub-rectangular mesh; branches broadly rounded with distinct carina which bears large, circular, widely spaced nodes; apertures in two rows with a third appearing before bifurcation; apertures medium-sized, circular, not stabilized, with operculum-like plate bearing a central boss; ornament of pustulose striations; zooecial bases elongate, irregularly pentagonal. Description. Zoartum: Gently to rapidly expanding fan-shaped fragments of unknown orientation; maximum radius 60mm. Obverse Surface: (a) Branches. Medium to wide (m.BW 0.40mm), frequently collapsed and distorted during preservation; branches sub-parallel to radiating in arrangement; branch cross-section rounded to triangular on either side of a sharply defined, broad carina; ornament of fine, wavy, pustulose striations. (b) Déssepiments. Medium width (m.DW 0.16mm), centrally straight, expanding only at branch junctions; ornament as on_ branches. (c) Fenestrules. Rectangular to sub-rectangular; regular mesh of medium length, medium width fenestrules (m.FL 1.63mm; m.FW 0.91mm). (d) Carina. Strong, massive, high carina varying from broad to sharp profile within one zoarium, the variation being a product of distortion during preservation; at bifurcation, carina swings on to one branch with a second, new carina commencing near a small node in the fork, or forming outside the third row of zooecia sometimes developed just before bifurcation. (e) Nodes. Large, circular nodes with elongate oval bases; distantly spaced (m. N-N 0.81 mm) in a single central row. (f) Zooeczal apertures. Medium size, circular (m.ZD 0.14mm); surrounded by a low, circular peristome, best Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 161 developed on the fenestrular rim; each bears an operculum-like plate with a central boss; apertures distantly spaced (m. Z-Z 0.42mm) with from three to five zooecia per fenestrule (m. Z/F 4.0) ; apertures in two unstabilized rows per branch with a third row up to Imm before branch bifurcation. (g) Addztzonal features. Hemispherical depressions (0.8-0.9mm diam.) occur on obverse surface near base of zoarium. Reverse surface. (a) Form. Rounded branches joined by medium-width dissepi- ments; branch width increases very close to bifurcation producing a distinctive tuning-fork shape; ornament of pustulose striations together with distally-directed irregular spines on a few branches. (b) Zooeczal bases. Elongate, irregularly pentagonal. Materzal. Holotype QUF24952 (Malchi Creek); Paratypes QUF24953 (Malchi Creek), QUF25006 (Por. 201-2, Parish Palen, Mt. Barney), SUF7432 (NULY) ; Others QGSF10903, 10904 (Malchi Creek), NEUF4705H, 4709A, 4718, 4739 (NUL454). Remarks. Deflation during preservation has caused much confusion with this species. The styles of deformation present are discussed with A. s. stroudenses (Campbell) . Campbell (1961) described two species A. malchi (Crockford) and A. cerva (Campbell) from Booral, N.S.W. and it is suggested in this paper that most of the differences between these two are a direct result of preservation. Distinguishing features listed by Campbell for A. cerva include large fenestrules, more zooecia per fenestrule, a more defined carina and peristomes (not recorded for A. malchz). The first two differences have been bridged in the present study by intermediate forms and the latter two are preservational aspects, rather than specific differences. Tall nodes recorded on A. cerva by Campbell have also been found in specimens of A. malchi. For these reasons, the two taxa have been contracted into A. malchi in the present description. A. malchi is also similar to A. macleayensss (Campbell, 1961), the major differences being largely that of size of fenestrules, branches and nodal spacing. Since the present study has failed to produce transitional material, these two taxa have been retained in their present form. Stratigraphy. A. malchi occurs in the Levipustula levis zone at Malchi Creek and Mt. Barney in Queensland and at several localities in the Stroud-Gloucester and Myall Synclines (including Booral) in New South Wales. Present information suggests that the species ranges throughout most of the brachiopod assemblage zone. Australofenestella macleayensis (Campbell, 1961) Fig. 5, 9-11 1961 F. crockfordae Campbell, pp.457-458, pl.59, figs. 2a-b. [non] 1960 F. (Mznilya) crockfordae Burckle, p.1088. 1962 F. macleayensis Campbell, p.48, pl.11, figs. lla-c. Revised diagnosis. Fenestrate with coarse, regular, sub-rectangular mesh; straight, wide, broadly rounded branches with a high, rounded, well-defined carina which bears distant nodes; two rows of unstabilized apertures per branch with increase to three rows prior to bifurcation; apertures medium-sized, erect with entire peristome within which there is an operculum-like plate with a central boss; branch ornament of fine pustulose striations; zooecial bases elongate, irregularly pentagonal. Description. Zoartum: Flat, slightly expanding fragment of unknown orientation; maximum radius 50mm. Obverse surface: (a) Branches. Straight, wide (m. BW 0.47mm), commonly deflated; profile broadly rounded with a sharply-defined central carina; ornament of fine pustulose striations. (b) Dzssepiments. Medium to Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 162 FENESTRATE BRYOZOANS wide (m. DW 0.18mm); centrally straight with gradual expansion to branch junction; ornament as for branches. (c) Fenestrules. Sub-rectangular, coarse, regular mesh; fenestrules long and wide (m. FL 2.62mm; m. FW 1.04mm). (d) Carina. High, prominent, well-rounded, medium to broad, being less than one third of the width of the branches; with deflation, carinae become much broader in profile; two carinae develop with the early appearance of a third row of zooecia before bifurcation. (e) Nodes. High, bladed nodes situated on the central carina; spacing very distant and irregular (m. N-N 1.10mm) ; nodes poorly preserved on holotype. (f) Zooeczal apertures. Medium-sized, circular (m. ZD 0.15mm), with slightly raised peristome producing erect, cup-shaped apertures; apertures display an operculum- like plate with a central boss; apertures not stabilized; widely spaced (m. Z-Z 0.43mm) with from three to six and one half zooecia per fenestrule (m. Z/F 6.1) being arranged in two rows with a third row appearing up to 1.6mm pre-bifurcation. Reverse surface. (a) Form. Rounded branches joined by rounded dissepiments less than half branch width; ornament of fine, close striae generally obscured by over- growth. (b) Zooeczal bases. Elongate, irregularly pentagonal. Materzal. Holotype NEUF5738 (NUL390 — Oaky Creek, Kempsey); Others NEUF4698, 4699 (NUL9 — Booral). Remarks. Campbell (1961) erected a new species A. crockfordae (Campbell) which is pre-occupied by F. (Minilya) crockfordae Burckle (1960) and therefore must be replaced. Campbell (1962) described a further new species A. macleayenszs which in the opinion of Wass (1968, pp. 83, 85) and the present writer is specifically identical with A. crockfordae (Campbell). The former name has therefore been selected for this taxon. Reasons for the contraction are based upon the following: Campbell (1962) noted that the two species were comparable, but decided that A. crockfordae (Campbell) was distinctive because of its large nodes and very wide carina. As A. macleayensis was described as being nodeless, this represented a significant difference. However, careful re-examination of the latter holotype reveals that it does have nodes which are difficult to observe because of their very poor preservation. When measured they have identical spacing with those of A. crockfordae (Campbell). Carinal differences between A. crockfordae (Campbell) and A. macleayensis are due to different modes of deformation during preservation as discussed in the remarks with A. stroudensis (Campbell). As a result, it becomes apparent that only one species is now required. Stratigraphy. This species is known only from the Levipustula levis zone in New South Wales. Australofenestella cincta (Crockford, 1949) Fig. 4, 10-11; Fig. 5, 1-3 1949 Fenestrellina cincta Crockford, p.425, text-fig. 8. 1961 Fenestella cf. cincta (Crockford) , Campbell, p.460. 1964 F. czncta (Crockford) , Maxwell, p.38, pl.13, fig. 1. 1972 F. (Bajoola) cincta (Crockford) , Fleming. p.4, pl.2, figs. 4-6. Revised diagnosis. Coarse fenestrate with very wide, rapidly-bifurcating branches forming a coarse, irregularly rectangular mesh; branches with triangular profile, bearing a strong, very high, nodeless carina; apertures in two rows with no pre- bifurcation increase; apertures large, circular, erect, and of cup-like form with the peristome being raised on the fenestrular margin; branch ornament of distinctive, pustulose, longitudinal striations; zooecial bases very elongate, irregularly pentagonal. Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 163 Description. Zoartum: Narrowly radiating to sub-parallel, mature fragments of unknown orientation; maximum radius 40mm; branch bifurcation frequent. Obverse surface: (a) Branches. Straight to slightly wavy, sub-parallel; very wide (m. BW 0.55mm); rapid spread of branches after bifurcation followed by sub-parallel development has produced a distinctive ‘tuning-fork’ arrangement; cross-section tri- angular to sub-rounded, tapering steeply upwards to a very prominent central carina; ornament of fine to coarse, longitudinal, pustulose, sinuous _ ribbing. (b) Dzssepements. Medium to wide (m. DW 0.19mm) ; centrally straight with minor expansion at branch junction; situated at or just below branch level; ornament of prominent ribbing which is continuous on to branch sides. (c) Fenestrules. Very coarse, irregular mesh; high frequency of bifurcation, and the growth of dissepiments not always perpendicular to branches, disrupts an otherwise rectangular outline; fenestrules of variable dimensions; usually very long and wide with some fenestrules being very short (m. FL 3.39mm, O.R. 1.59-6.9mm; m. FW 1.23mm, O.R. 0.7- 2.0mm). (d) Carina. Acutely triangular, very high, slightly sinuous median carina; considerably steeper than branch profile from which it is separated by a marked change of slope; carina continuous on one branch at bifurcation, showing a tem- porary loss of profile adjacent to the zooecial aperture in the fork, new carina on the other branch forms up from one of a group of prominent surface ribs located on the side of the branch, there being no obvious connection back to the primary carina. _(e) Nodes. Absent. (f) Zooeczal apertures. Large, circular (m. ZD 0.19mm) surrounded by an entire, strongly developed, exserted peristome; apertural profile low, adjacent to carina, but with maximum elevation on the fenestrular margin giving the aperture a cup-like form on the obverse slope of the branch; zooecia arranged in two rows with no pre-bifurcation increase except for one aperture being located on the fork at each branch division; apertures alternate in position in adjoining rows being not stabilized with respect to the dissepiments and displaying very slight fenestrular indentation; apertures closed by operculum-like plates, each of which bears a variably-placed, spine-like projection; apertures very widely spaced (m. Z-Z 0.49mm) with from 3 to 12 zooecia per fenestrule (m. Z/F 7.0). (g) Additional features. Occasional resorbed apertures are surrounded by a semi-circular pit located on the branch between the carina and a narrow semi-circular ridge which extends beyond the edge of the branch (0.36-0.42mm long by 0.24-3mm wide). The aperture in the base of the pit has a very reduced peristome but still has the opercular plate present. One aperture was observed to have a small spherical sac obscuring it. An ovicellular function has been postulated. Reverse surface: (a) Form. Rounded branches joined by small, medium-width level dissepiments; branch surface longitudinally ornamented with fine, pustulose ribbing ; similar, but coarser, ribbing occurs on the dissepiments. (b) Zooeczal bases. Very elongate, irregularly pentagonal. Material. Holotype lost; Neotype (Fleming, 1972) QGSF10898 (Neerkol Fm., Ridgelands 1 mile map ref. 265791); Others QGSF10891, 10897 (topotypes) ; NEUF4715E, 4719, 4700A (NUL9); NUF2377 (NUL472); numerous additional localities in the Stroud-Gloucester-Bulahdelah region, N.S.W. Remarks. This most distinctive, very large species exhibits strong development of the operculum-like plate, with its variably placed boss-like projection, which closes the external aperture. The function of the projection on the plate cannot be determined from the external moulds available. It is considered to represent either a spine on the surface or a tube-like extension through the plate. A. cincta differs from all other species described in this paper in possessing a very high, bladed, nodeless carina which separates the two rows of apertures on each Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 164 FENESTRATE BRYOZOANS branch. In this respect, the inclusion of A. czncta in this paper is somewhat anomalous except for the apertural development which is considered to be its most significant feature. Fleming (1966) placed A. cencta in his subgenus Fenestella (Bajoola), which was defined to include species of Fenestella with strong nodeless carinae. Detailed com- parison of the type species F. (Bajoola) capellae Fleming with the present species reveals a number of dissimilarities which would suggest that the grouping is possibly inappropriate. Significant differences in the structure of the carina and of the zooecial apertures and chambers, form the basis upon which the grouping has not been per- petuated. Other Early Carboniferous species (F. rowchels Crockford, F. propinqua de Koninck and F. brownei Roberts) have much more in common with the type species and may possibly be grouped with it at some future date. A. cincta remains a solitary form without obvious affinity to any other species known to the writer. No useful comparisons can be offered from the literature at this stage. Stratigraphy. A. cincta is widely found associated with the Lewpustula levis zone. Its occurrence has been reported in Queensland from the Stanwell district (Crockford, 1949; Fleming, 1972) and the Yarrol district (Maxwell, 1964) in beds equated to various levels in the Neerkol Formation. New South Wales material has been described from Booral (Campbell, 1961) and has been found in the present study in a number of other localities in the Stroud-Gloucester-Bulahdelah region. Australofenestella(?) keepitensis sp.nov. Fig. 4, 1-3 1963 Fenestella sp.1. Campbell & Engel, pp.67-68, pl.1, figs. 4-5. Diagnosis. Coarse, irregular fenestrate with wide, straight to slightly wavy, non- carinate branches forming. a sub-oval to sub-rectangular mesh; nodes very large, widely spaced, irregularly distributed; apertures moderately large, not stabilized, situated low on branch sides; zooecia in two rows with a third, intermittently developed, central row of apertural-sized pits which are developed unrelated to branch bifurcations; zooecial bases large irregularly pentagonal in lateral rows and rhomboidal in the central row when developed. Description. Zoartum: Expanding fragment of unknown orientation; maximum radius 65mm. Obverse surface: (a) Branches. Wide (m. BW 0.46mm), straight to slightly wavy with gently radiating arrangement; branch cross section rounded; ornament of fine pustulose striations obscured by secondary overgrowth. (b) Disseprments. Wide (m. DW 0.42mm); branch junctions rounded to sub- rounded; situated below branch level; no ornament observed. (c) Fenestrules. Sub- rectangular to sub-oval; irregular mesh of long, wide fenestrules (m. FL 2.61mm; m. FW 1.10mm). (d) Carina. Absent; some low ridging occurs between nodes. (e) Nodes. Very large, prominent oval-based, round nodes (basal diameter 0.25- 0.4mm); medium to distant spacing (m. N-N 0.49mm); node arrangement extremely irregular, varying from a central row, to a zigzag pattern, to a few cases of two nodes being placed alongside each other; nodal variation is irregular without relationship to branch bifurcations. (f) Zooecial apertures. Medium to large, circular (m. ZD 0.16mm) ; surrounded by low peristome, best developed on the fenestrular rim; apertures in two unstabilized rows situated low on branch sides where the apertures face into the fenestrule; apertures distantly spaced (m. Z% 0.47mm) with from five to six zooecia per fenestrule (m. Z/F 5.6) ; irregularly developed short rows of aperture-like pits occur along the centre line of some branches. Development is unrelated to branch bifurcations but the branches show some thickening in the region Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 165 of these extra rows. The pits are located between nodes and represent the brief, inter- mittent development of a third row of apertures. Reverse surface: (a) Form. Irregular branches, varying from straight to zigzag in an anastomosing format; dissepiments equal in dimension to, and level with, branches; surface smooth. (b) Zooeczal bases. Large, irregular, pentagonal in lateral rows and rhomboidal where a third row is developed. Materzal. Holotype NEUF7466A/B (NEUL318 — Swaines Gully, Werrie Syncline) . Remarks. Evidence that the irregular central occurrence of small pits represents additional apertural development comes from the observation of three rows of zooecial chambers in some parts of the zoarium. Peristomal development is very weak on these central apertures. The distribution of nodes and the erratic development of a third central row of apertures together with the absence of a central carina make it very unlikely that this species belongs to Fenestella Lonsdale. Although zigzag nodes are a feature of the subgenus F. (Mznzlya) Crockford, the nature of their development in the present species prevents any viable comparison being made. No other species in the literature at present available can be usefully compared with A. (?) keepztenszs which can only be dubiously assigned to this genus. Stratigraphy. The species is known only from the Tulcumba Sandstone where it is associated with a Spzrifer sol assemblage of early Tournaisian age. COMPARATIVE GROUPING OF SPECIES Tables 1 and 2 detail the descriptive and statistical aspects of all the species/ subspecies described in this paper. Multi-rowed species. Stable, unifying features which justify the grouping of the multi- rowed species adopted in this paper include: (a) Apertures of large size, and medium to distant spacing, all of which bear a wide, low, peristomal collar which may be entire, or horseshoe-shaped with a proximal opening. All apertures bear a transverse plate or operculum and this plate carries a raised spine or boss-like projection, the position of which is usually central, but can be eccentrically placed; (b) Straight, wide to very wide branches of oval to flattened cross-section. Most species bear moderate to very strong development of distinctive, pustulose ribbing. All species lack a central carina having at most a low, internodal rise of very poor development; (c) Nodes are generally absent from the group but where developed are located on the proximo-central rim of an aperture in any zooecial row. In two species (Australo- polypora rawdonensis and Australopolypora altinodosa), the obverse surface bears large, widely-spaced spines of presumed attachment significance because of their robust construction and lack of outward termination. Some species exhibit a greater degree of similarity than others and the following discussion relates to these forms: (1) Australopolypora rawdonensis and Australopolypora palenensis. ‘hese two species have the coarsest mesh of the multi-rowed species with a fenestrule length close to 3mm and a width of about 1.3mm. From the reverse, the form, ornament and zooecial bases are very similar. The major distinction lies in the position, form, and spacing of the zooecial apertures. Australopolypora rawdonensis has horseshoe-shaped peristomes surrounding oval apertures which are placed in a fan-like arrangement. Central apertures have their long axis parallel to the branch length whereas the side apertures are inclined at 45 degrees to'the length. By contrast, Australopolypora palenensis has apertures which are distally inclined to the branch surface, having the proximal Proc. Linn. Soc. N.S.W., 103 (3), (1978) 1979 166 FENESTRATE BRYOZOANS TABLE 1 A descriptive comparison of the important morphological features of the fenestrate mesh of all species of Australopolypora gen.nov. and Australofenestella gen.nov. described in this paper. En en BY ay ae ne 2}Fn0|B/10 2/5 |Z/F Soe ; 1:75 0-93 0-46 0-17 0-20 0-49 0-90 20 | 5-7 |10-7}10-2 2 R|1-40- 2:10 | 0-66-1-10 }0-40-054/0-14- 0-24} 0-18-0-22/0:40-060 0:42-1:26 6-2 }10-1 }12-4 A O.R11-14 - 2:30} 0-64 -1:80}0-36 - 1-16 |042 - 0-52} 0-16-0286 10-30-0 62 J0-72-1:20 3 1:56 1-14 0-86 0-74 0-23 0-41 6-4 | 8-7}12-2| 3-8 scalpta AUSTRALOPOLYPORA 1:20-1:84 |0-88-1-60)0-60-1-16 ]0-52-1-12} 0:20-0:24/0-34-056 D ° ; 0-50 é é : rare {140 19-1]3-1 keppelensis : . 0-057 ; : keppelensis 5 ; Ske oh : 0-64 e he ° : present |}40 |12:0 |15-6]19-3] 3-2 Keppelensis 0:072 | 0-058 | 0- parvula ae 0-68-0395 |050-0-84 |0-32:0-60 |0:20-0:30]0-12-0 18 0:22-0:32 1-49 0-72 0-33 0-17 0-15 0-29 0-71 3-9]17-2)5-1 brooker! : O.R.|1:10-1-72 |0-60-0-9 0} 0:28-0:38 }0-12-0-20,0-14-9-18/0-26-0-32)054-0590 stroudensis 1-37 0:72 0-29 0:08 0:17 0:30 0-44 ||20 | 7:3 |13-9 16-8 | 4-6 MINUET |g R 1956-494 |0:50-0:92|0-26-0-34|0-04-0-12|0-14-0-18 |0-26-034]0-28-0-60 : 4 : 5-2 |10-3/11-8 [4-5 stroudensis stroudenss ; ; 0 0 ¢ : : c 6-2 }11-1112-2] 4-0 malchi : ; : “045 : . : 112-2: i 7 24-0: : ‘ 10-0- 5 s é . 5 ° cd é = : 3-8 11°6 macleayensis a F | : E | : 2-61 0-46 0-42 : : e 20| 3:8 | 9-1 | 10:8} 5-6 (?)keepitensis R| 2-16 -3-28) 0-76 -1-4 8) O- 36- 054 0:28-0-5 J 0:14-0:18) 0:38-0:54 0-36-0:80 rim of the peristome level with the surface and the distal rim depressed below branch level. (2) Australopolypora altinodosa and Australopolypora neerkolensis. These two species have very close mesh dimensions. A. altinodosa has slightly narrower branches in keeping with its more extensive development of two rows of zooecia. AUSTRALOFENESTELLA Proc. LINN. Soc. N.S.W., 103 (3), (1978) 1979 B.A. ENGEL 167 bear large attachment spines as noted above. In most respects, these species strong affinity with the number of zooecial rows being the principal distinc- Since A. neerkolensts has not been found at the one locality where A. 09 =a Q »dosa occurs, it is possible that the latter is just a local variant of A. ; a0 zs ‘olenszs at Booral, N.S.W. Lack of further material precludes a definite | < a mM ton. : : uv gc S remaining taxa Australopolypora scalpta and Australopolypora keppelensis c S sla are distinctive in their own right and exhibit no obvious grouping < ® Ze res. @ + w =| @O se ee’ ie)) = TABLE 2 il summary of the mesh dimensions of all species of Australopolypora — Australofenestella n this paper. Species known only by a single specimen have only their means and observed ranges ‘or explanations of abbreviations see Engel (1975, p.577). Zooecia per enestrula Zooecial Ro post/norm] pre -al | bif Nodal |Apertuml Spacing] Size & & Size [Spacing distant Special Fenestrules Eaatines ies sub-oval to rectangular, long; wide sub-oval to rectangular. very wide; straight, | 2-3/3-4/4-5 CHASIS i=) ie) ~ Ni — ° 38) RZ) o,) OF 1 — OT ~ $11e} soy soz Bojode uopipa ouy peey ey, sepun pejujsd uaeq sey 1 ojqey jo uey4ye medium length, medium width lensis parr short length, FVuia medium width regular|round-oval medium,| narrow, blunt, Australasian Rodents ROBERT DOMROW Domrow, R. Some dermanyssid mites (Acari), mostly from Australasian rodents. Proc. Linn. Soc. N.S.W., 103 (4), (1978), 1979: 189-208. Results are presented from three sponsored expeditions to collect parasitic mites from mammals, mostly rodents, in remote parts of Australasia. Three new species of Laelaps are described from native-mice (Pseudomys spp.) in Western Australia: L. janalis from P. occzdentalis, L. bycalza from P. albocinereus, and L. lybacia from P. praeconis. Additional data, including new hosts and localities, and previously unknown males and immature stages, are given for a further 22 species in nine genera: Mesolaelaps (3), Haemolaelaps (2), Peramelaelaps (1), Laelaps (9), Eulaelaps (1), Echinonyssus (1), Trichosurolaelaps (2), Ornithonyssus (2) and Halarachne (1). A small group of intractable specimens of Laelaps with both setae on coxa I simple is still under study. R. Domrow, Queensland Institute of Medical Research, Bramston Tce, Herston, Australia 4006; manuscript received 10 January 1978, accepted in revised form 23 August 1978. This paper assembles results from three Australian Biological Resources Study grants, gratefully acknowledged — one to Dr C. H. S. Watts, Institute for Medical and Veterinary Research, Adelaide, for a circum-Australian sampling of rats (Robinson et al., 1978) and two to me to collect ectoparasites in remote areas of Queensland. For brevity, collectors are omitted from Dr Watts’ offering (with map coordinates), but I hasten to add that they were A. C. & J. F. Robinson. Their material will be deposited in the Australian National Insect Collection, CSIRO, _ Canberra. The data for my contributions are: Bamaga, near Cape York, iii.1975, R. Domrow and J. S. Welch; and Kowanyama (formerly Mitchell River Mission) , east coast of Gulf of Carpentaria, vi.1976, R. Domrow. These two collections will be divided between the Queensland Museum, Brisbane, and my institute (QIMR). Lastly, a few recent accessions in QIMR are added, with data in full. I thank all concerned. Terminology is largely after Evans and Till (1965), with tarsi II-IV after Evans (1969) . Hosts are after Ride (1970), with subspecies for rats supplied after Taylor and Horner (1973). Mesolaelaps australiensis (Hirst) Laelaps (Mesolaelaps) australiens’s Hirst, 1926, Proc. zool. Soc. Lond., 1926: 840. Material. Three 9°, Rattus fuscipes fuscipes (Waterhouse) (three rats), 22 km NE Jurien (30°8’, 115°8’), W.A., 12-13.iv.1975; 10 29, R. f. coracius Thomas (three rats), 41 km SE Cairns (17°15', 145°56’), Qd, 16.xi.1974; seven 99, R. leucopus leucopus (Gray), Bamaga; one Q, R. lutreolus lutreolus (Gray), 20 km SW Port Macquarie (31°37, 152°50), N.S.W., 2.i1.1975; 14 992, R. sordidus sordidus (Gould) (three rats) , Iron Range, 23 km S Portland Roads (12°47', 143°18') , Qd, 2- 3.xi.1974; two 99, R. s. sordidus (two rats), 37 km S Cooktown (15°48', 145°14’), Qd, 9.xi.1974; 10 99, R. s. sordidus (four rats), 16 km S Cairns (17°4’, 145°47’), Qd, 14.xi.1974; eleven 92, one deutonymph (hereafter abbreviated as dn), R. s. sordidus (four rats), 11 km NE Atherton (17°12’, 145°33’), Qd, 22.xi.1974; one , Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 190 SOME DERMANYSSID MITES (ACARI) R. s. villosissimus (Waite), 2 km NE Mount Isa (20°38’, 139°30’), Qd, 19.x.1974; one 9, R. s. villosissemus, 32 km W Windorah (25°20’, 142°18’), Od, 15.x.1974; one 2, R. tunneyt tunney: (Thomas), 14 km S Nourlangie Camp (12°54’, 132°39’), N.T., 18.vi.1975; one 9, R. t. culmorum (Thomas and Dollman), 58 km N Maryborough (25°6', 152°33'), Qd, 14.1.1975; one 9, Hydromys chrysogaster Geoffroy, Maslin Creek, Atherton (17°15, 145°29'), Qd, 24.xi.1974; one 2, Melomys littoralis (Lénnberg), 11 km NE Atherton (17°12’, 145°33’), Qd, 22.xi.1974; one 9, M. littoralés, 9km SE Dunwich, Stradbroke Island (27°32’, 153°30'), Qd, 19.1.1975. Deutonymph. Capitulum 355 um long, as in M. antzpodianus (Hirst) (see Domrow, 1977), but setae c and h; well exceeding sides of basis. Epistome an equilateral triangle, free sides ever so slightly convex, with weak denticulations and broad, but fine, submarginal dendritic pattern. Chelicerae stronger, 248 um long, with digits more elongate, occupying one-quarter of total length.* Idiosoma 780 yum long, 530 wm wide (somewhat engorged). Dorsal shield 715x335 um, holotrichous, differing from that of M. antzpodzanus only in that setae are slightly more elongate. Most of Z and S setae broken, but Z; certainly elongate. Setae on cuticle longer and more widely spaced than in M. antipodzanus (as is case in adults of two species) . Venter as in M. antzpodzanus, but genital portion of sternogenital shield tapering very sharply behind metasternal setae, parallel-sided, only half as wide as space between genital setae; setae on cuticle again longer and more widely spaced. Postanal seta broken off, but its insertion well exceeded by adanals. At least one small metapodal shield on each side. Legs holotrichous except for one additional v seta on tibia I (2-6/4-2) as in M. antipodianus (¢ also showing this additional seta). Longest seta on dorsum of tarsus IV 140 um. Notes. M. australzensis is widespread in Australia, showing a low level of host- specificity (Domrow, 1961, 1962a, 1967). At a subspecific level, R. f. fusczpes, R. I. leucopus, R. s. villosissimus, R. t. tunney and R. t. culmorum are new host-records, R. t. tunneyt extending the range of this mite into the Northern Territory. Extra- Australian records (New Guinea, New Zealand and Kermadec Islands) were summarized by Tenorio and Radovsky (1974). Mesolaelaps bandicoota (Womersley) Hypoaspis bandicoota Womersley, 1956, Linn. Soc. J., Zool., 42: 573. Material. Two 99, Rattus fuscipes assimilis (Gould), Mount Stanley, 38 km E Kingaroy (26°30, 152°13’), Qd, 16.1.1975; four 99, R. f. ass¢miles (two rats) , 56 km SE Canberra (35°41, 149°32’), N.S.W., 14.11.1975; one 2, R. f. assimilis, 20 km NE Mallacoota (37°27, 149°57'), N.S.W., 17.11.1975; two 99; R. lutreolus lutreolus, 6 km SW Bemm River (37°47', 148°54’), Vic., 21.11.1975. Notes. This material confirms earlier records (Domrow, 1963, 1973). All specimens show the anterior seta on coxae II-III normal (unexpanded) and the dorsal shield holotrichous, except one female from near Canberra with three setae at J, (Domrow, 1977). Mesolaelaps anomalus (Hirst) Laelaps (Mesolaelaps) anomalus Hirst, 1926, Proc. zool. Soc. Lond., 1926: 840. Material. Two 29, Isoodon macrourus (Gould) , Bamaga. Notes. This material confirms previous records (Domrow, 1962a, 1967). *Proportions, rather than absolute measurements, seem important with single deutonymphal specimens, since prefemales differ from premales only in their larger size. ~ Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 191 Figs 1-3. Haemolaelaps domrouwt. 1-2. Idiosoma in dorsal and ventral views, pn. 3. Epistome, 9. Note: the subdivisions on scales with Figs 1-23 represent 100 pm. Haemolaelaps domrowt Womersley (Figs 1-3) Haemolaelaps domrowt Womersley, 1958, Proc. Linn. Soc. N.S.W., 82: 301. Material. Six 92, one d, two protonymphs (hereafter abbreviated pn), Isoodon macrourus, Bamaga. Female. Epistome slightly shorter and more rounded than in H. flagellatus Womersley (see Domrow, 1977), with fewer, slightly stronger denticles and submarginal dendritic pattern as in H. calypso Domrow, 1966. Protonymph. Basis capituli slightly longer than wide, with setae c slightly more than half as long as interspace, exceeding sides of basis; deutosternum with at least five rows of denticles, mostly double. Hypostome with setae h3>h,>h,; h, almost one and a half times as long as interspace; hf; slightly more than one and a half times as long as interspace, slightly exceeding sides of basis. Labial cornicles pale, but with opposed tips. Epistome anticipating that of 9. Palpal setation (trochanter-tibia) holotrichous, i.e. 1.4.5.12 (including two dorsodistal rods) ; seta al, on genu spatulate; tarsus with one of three v elongate; claw bifid. Chelicerae as in Q, i.e. fixed digit straight, weak, edentate, with elongate pilus dentilis (small dorsal seta present, but pores not detected) ; movable digit stronger, with two small teeth near incurved tip. Idiosoma 410-455 ym long, 260-300 um wide. Surface of dorsal shields marked by paired muscle insertions and reticulate. Podonotal shield with three extremely shallow lobes posteriorly; podonotum holotrichous, with 16 pairs of setae (eleven on shield, five on cuticle). Pygidial shield semicircular, with pair of very distinct pores in anterolateral angles in front of setae S,. Opisthonotum holotrichous, with 14 pairs of setae (eight on shield, six on cuticle). All setae subequal except for slightly smaller 7, and J;, and much longer Z, (lattermost sinuous, but this may be artifact of mounting) . Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 192 SOME DERMANYSSID MITES (ACARI) Tritosternal base with a few distinct barbs laterally as in adult; laciniae ciliated. Sternal shield elongate, with some sign of reticulation; anterior margin weak, preceded by narrow zone of striae; lateral margins straight between insertions of setae; posterior margin triangular, with indication of short backward extension; with usual three pairs of setae and two pairs of pores. Genital complex represented only by pair of small setae. Anal shield appearing slightly foreshortened posteriorly, with postanal seta twice as long as adanals. Ventral cuticle with three pairs of setae in front of, and one pair flanking, anal shield. Peritremes short, not reaching forward beyond articulations on coxae III; peritrematal shields, if present, completely lateral and out of sight. Legs largely folded under, but setation seen to be holotrichous. No d seta on femora-genua I-IT unduly lengthened. Notes. Immatures of this species were previously undescribed. The material confirms earlier records (Domrow, 1962a, 1967). Haemolaelaps penelope Domrow Haemolaelaps penelope Domrow, 1964, Proc. Linn. Soc. N.S.W., 89: 156. Material. One 2, Trichosurus caninus (Ogilby) , Clouds Creek, N.S.W., 1x.1977, J. H. Arundel. Notes. The only previous record of this species was from S.E. Queensland. Peramelaelaps bandicoota Womersley (Figs 4-7) Peramelaelaps bandicoota Womersley, 1956, Linn. Soc. J., Zool., 42: 574. Materzal. Eleven 9, five dd’, two dn, one pn, Isoodon macrourus, Bamaga. Female. Basis capituli longer than wide, with setae c of moderate length, about two- fifths as long as interspace, just reaching sides of basis; deutosternum with six rows of one to three denticles. Hypostome with setae h3;>h,>h,; h, slightly longer than interspace; h3 one and a quarter times longer than interspace, well exceeding sides of basis. Labial cornicles pale, but with opposed tips. Labrum slowly tapering, pointed, spiculate. Epistome soft and diaphanous, with smooth margin and submarginal trace of dendritic pattern reaching to midlength, about twice as long as basal width and roundly pointed, reaching just beyond distal margins of palpal femora. Palpal setation (trochanter-tibia) holotrichous, i.e. 2.5.6.14 (including two dorsodistal rods) ; seta al, on genu spatulate; tarsus with one of three v elongate; claw bifid. Chelicerae with fixed digit pale, straight, edentate (dorsal seta and pores not detected) ; movable digit well sclerotized, with two teeth more distinct than originally figured. Dorsal shield with surface weakly marked by paired muscle insertions, not reticulate except for vertical, humeral and lateral band that narrows and disappears posteriorly; pores probably more numerous than figured; setation holotrichous (22 pairs of podonotal setae, 17 pairs of opisthonotals — S, lacking on one side of specimen figured) . Tritosternal base with fine, soft fringe laterally, cf. H. domrowz above. Leg setation holotrichous. No d seta on femora-genua I-II unduly lengthened. Male. Capitulum as in 9 except for shorter setae h; (about half as long as interspace, falling short of sides of basis) and spermatodactyl. Latter with spermatophore-carrier stout, abruptly and shortly bent apically around tip of somewhat reduced movable digit. Idiosoma 380-395 wm long, 210-240 um wide. Dorsum as in @, but almost completely covered by dorsal shield and with only about two pairs of setae on cuticle. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 193 Figs 4-7. Peramelaelaps bandicoota. 4-5. Idiosoma in dorsal and ventral views, 9. 6-7. Idiosoma in dorsal and ventral views, d. Sternal portion of holoventral shield as in 2, but more arched anteriorly to accept genital aperture; ventral portion expanded and normally sharply angulate behind coxae IV, with four (at times five) pairs of usurped ventral setae. Ventral cuticle with about eight pairs of setae of increasing length posteriorly. Metapodal shields at times insensibly fused into holoventral shield. Peritremes a little shorter than in 9. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 194 SOME DERMANYSSID MITES (ACARI) Legs as in 9, but spurs on coxae simpler and some ventral setae on II strengthened: av on femur, av and pv on genu (to some extent) and tibia, av,_; and pv2 On tarsus. Deutonymph. Capitulum not clear in either specimen, but essentially predicting that Gree Idiosoma 475 um long, 265 wm wide (enclosing developing @) ; 405 ym long, 235 um wide (enclosing developing 6). Dorsal shield with lateral incisions narrow, reaching slightly past midpoint between setae J, and Z,; setation of prefemale obscured by that of developing adult, but shield of premale torn free of developing adult and clearly holotrichous (22 pairs of podonotal setae, 17 pairs of opisthonotals) . Sternogenital shield with four pairs of setae and three pairs of pores; ligulate posteriorly, leaving genital setae and pores free in cuticle. Remainder of venter not clear due to doubling, but peritremes elongate. Leg setation not clear due to doubling, but probably holotrichous. Spurs on coxae simple. Protonymph. Capitulum predicting that of 2; palpal setation not clear because of doubling, but probably holotrichous. Idiosoma 385 ym long, 210 um wide (enclosing developing deutonymph). Podonotal shield lightly trilobed posteriorly, median lobe strongest. Pygidial shield almost straight anteriorly, but with slight median prominence; with pair of very distinct pores in front of setae S, as in H. domrowz above. Setation not clear because of doubling, but probably holotrichous. Sternal shield with three pairs of setae and two pairs of pores, but outline not clear. Genital complex represented at least by a pair of small setae. Ventral cuticle with three pairs of setae in front of, and one pair flanking, anal shield. Peritremes abbreviated. Leg setation not clear because of doubling, but probably holotrichous. Notes. The above description of the female of this species is only so full as to complete Womersley’s text; the male and immatures were previously undescribed. These specimens confirm earlier records (Domrow, 1962a, 1967). Laelaps southcott? Domrow Laelaps southcottt Domrow, 1958, Proc. Linn. Soc. N.S.W., 82: 364. Material. Six 22, two pn, Uromys caudimaculatus (Krefft) (two rats), Iron Range, 19 km SW Portland Roads (12°43, 143°17’), Qd, 2.xi.1974; four 29, U. caudimaculatus (two rats), 40 km SE Cairns (17°15’, 145°56’), Qd, 15-16.xi.1974; three pn, U. caudimaculatus, 19 km SE Atherton (17°25', 145°31°) , Qd, 25.xi.1974. Notes. This material confirms the original record. Laelaps sp. Material. Many specimens, Rattus leucopus leucopus, Bamaga; ten 2@, four dn, three pn, R. /. leucopus (three rats) , Iron Range, 19 km SW Portland Roads (12°43’, 143°17'), Qd, 3.xi.1974; one Y, one dn, R. 1. leucopus (two rats) , Iron Range, 20 km SW Portland Roads (12°44’, 143°16'), Qd, 31.x.1974; one dn, one pn, R. 1. leucopus, Iron Range, 27 km S Portland Roads (12°49’, 143°18’), Qd, 2.xi.1974; three 92, one dG, three dn, R. sordidus sordidus (two rats), Iron Range, 23 km S Portland Roads (12°47’, 143°18’) , Qd, 2.xi.1974; two 99, R. tunneyi culmorum, 58 km N Rockhampton (22°52’, 150°41’) , Qd, 5.i1.1975. Notes. These difficult specimens, with both setae on coxa IJ simple, will be treated later. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 195 Laelaps nuttalli Hirst Laelaps nuttalli Hirst, 1915, Bull. entomol. Res., 6: 183. Material. Two 99, Rattus rattus (Linnaeus), Mamara, Guadalcanal, Solomon Islands, 12.xi.1975, J. A. R. Miles; one 9, R. rattus, Mendana, Ndende, Santa Cruz Group, Solomon Islands, 31.x.1975, J.A.R.M.; two 29, one dn, R. rattus, 11 km NE Atherton (17°12', 145°33’), Qd, 22.xi.1974; one d, R. exulans (Peale), Honiara, Guadalcanal, 28.x.1975, J.A.R.M.; three 99, R. exulans, Pamua, Makira (= San Cristobal) , Solomon Islands, 8.xi.1975, J.A.R.M.; four 99, one d, R. exulans, Kira Kira West, Makira, 9.xi.1975, J.A.R.M.; one 9, R. exulans, Mendana, 31.x.1975, J.A.R.M.; one @, R. exulans, Graciosa Bay, Ndende, 1.xi.1975, J.A.R.M.; nine QQ, two 6d, R. exulans, Onetar, Gaua, Banks Islands, New Hebrides, 23.x.1975, J.A.R.M.; five QQ, R. lutreolus lutreolus (two rats), 20 km SW Port Macquarie (31°37', 152°50°), N.S.W., 1.111975; one 2, R. sordidus sordidus, Kowanyama; six 292, R. s. sordidus (three rats), Iron Range, 23 km S Portland Roads (12°47’, 143°18), Qd, 2.xi.1974; three 99, two dd, four dn, one pn, R. s. sordédus (three rats), 16 km S Cairns (17°4', 145°47’), Qd, 14.xi.1974; three 29, one d, one dn, R. s. sordidus (three rats), 11 km NE Atherton (17°12’, 145°33’), Qd, 22.xi.1974; three 29, R. s. villostssimus, 56 km SE Boulia (15°22’, 140°), Qd, 18.x.1974; five 92, one 6, R. s. villostssemus, 77 km SE Boulia (15°35’, 140°8’), Qd, 17.x.1974; two 99, one 6, R. s. villostssemus, 2 km NE Mount Isa (20°38’, 139°30’), Qd, 19.x.1974; eight 22, six dd, seven dn, one pn, R. s. villosissimus (two rats), 33 km SE Richmond (20°49', 143°28'), Qd, 20.x.1974; 12 99, one dG, four dn, one pn, R. s. villostssimus (three rats), 32 km W Windorah (25°20’, 142°18’), Qd, 15.x.1974; 21 99, one dn, R. s. collettt (Thomas) (eight rats) , South Alligator River, 175 km E Darwin (12°42’, 132°32'), N.T., 7-8.vi.1975; six 99, two dn, one pn, R. s. colletté (four rats), Leanyer Swamp, 15 km E Darwin (12°23’, 130°56’), N.T., 11.vi.1975; two 99, one 6, one dn, R. tunney? tunneyi (two rats), 7 km SE Nourlangie Camp (12°49’, 132°42'), N.T., 13.vi.1975; one 9, Pseudomys gracilicaudatus (Gould), 98 km NW Bundaberg (24°31, 151°28’), Qd, 10.i.1975; four 99, one d, Melomys cervinipes (Gould) (two rats) , 32 km § Cooktown (15°45’, 145°18’), Qd, 7.xi.1974; one 9, M. cervinipes, 40 km SE Cairns (17°15', 145°56'), Qd, 15.xi.1974; one 2, M. littoralis, 22 km S Cooktown (15°39’, 145°13’), Qd, 7.xi.1974; one 2, M. littoralis, Leanyer Swamp, 15 km NE Darwin (12°22’, 130°56’), N.T., 11.vi.1975. Notes. All previous Australian records of this cosmopolitan parasite of small rodents were from Queensland (e.g. Domrow, 1958, 1962a) . Its range is now extended to the Northern Territory and New South Wales. At a subspecific level, R. 1. lutreolus, R. s. collett? and R. t. tunneyi are new host-records. Two recent references of a wider (Pacific) interest are Mitchell (1964) and Marshall (1976) . Laelaps assimilts Womersley Laelaps assimilis Womersley, 1956, Linn. Soc. J., Zool., 42: 557. Material. One ¢, Rattus rattus, 50 km NE Newcastle (32°39', 152°9’), N.S.W., 4.i1.1975; five 92, two dd, R. fuscipes assimilis (three rats) , Mount Stanley, 38 km E Kingaroy (26°30’, 152°13’), Qd, 16.i1.1975; three 92, two dd, one dn, three pn, one larva (hereafter abbreviated 1), R. f. ass¢mzlis (three rats), 20 km NE Mallacoota (37°27, 149°57), N.S.W., 16-17.ii1.1975; two 99, R. f. ass¢milis, 5 km SW Bemm River (37°47, 148°55') , Vic., 19.11.1975. Notes. This material confirms the original record. The specimen from R. rattus is a straggler. Laelaps wasselli Domrow Laelaps wasselli Domrow, 1958, Proc. Linn. Soc. N.S.W., 82: 363. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 196 SOME DERMANYSSID MITES (ACARI) Material. Three 92, Hydromys chrysogaster, Kowanyama; three 99, H. chrysogaster, Iron Range, 26 km SW Portland Roads (12°49’, 143°18'), Qd, 1.xi.1974; one pn, H. chrysogaster, 29 km SE Innisfail (17°46, 146°7'), Qd, 4.x11.1974. Notes. This material confirms the original record. Laelaps echidninus Berlese Laelaps (Iphis) echidninus Berlese, 1887, Acari, Myriapoda et Scorpiones hucusque in Italia reperta. Patavii. Fasc. 39, No. 1. Material. Eight 99, Rattus exulans, Mendana, Ndende, Santa Cruz Group, Solomon Islands, 31.x.1975, J.A.R.M.; one &, R. exulans, Loh, Torres Islands, New Hebrides, 16.x.1975, J.A.R.M.; five 92, one d, R. fuscipes assimilis (two rats), 56 km SE Canberra (35°41, 149°32’), N.S.W., 14.11.1975; three 99, R. f. assemilis, Wragge Creek, Kosciusko National Park (36°23, 148°28'), N.S.W., 11.11.1975; two QQ, one dn, R. f. assimilis (three rats), 20 km NE Mallacoota (37°27, 149°57’), N.S.W., 17.11.1975; one 9, R. f. ass¢milis, 6 km SW Bemm River (37°47, 148°54’), Vic., 20.ii.1975; many specimens, R. leucopus leucopus, Bamaga; 29 99, R. l. leucopus (three rats) , Iron Range, 19 km SW Portland Roads (12°43’, 143°17’), Qd, 1-3.xi.1974; two 99, R. l. leucopus, Iron Range, 26 km SW Portland Roads (12°44’, 143°14'), Qd, 31.x.1974; one pn, R. I. leucopus, Iron Range, 20 km SW Portland Roads (12°44’, 143°16’), Qd, 31.x.1974; nine 99, R. l. leucopus, Iron Range, 24 km S Portland Roads (12°47, 145°18’), Qd, 31.x.1974; one 2, R. 1. leucopus, Iron Range, 27 km S Portland Roads (12°49, 143°18), Qd, 2.xi.1974; 2199, R. 1. cooktownensis Tate (five rats), 32 km S Cooktown (15°45', 145°18), Qd, 7- 9.xi.1974; many 22, R. 1. cooktownensis, Mossman, Qd, vi.1970 and v.1971, R. Domrow and R. W. Campbell. Notes. At a subspecific level, R. 1. leucopus and R. |. cooktownensts are new host- records. Campbell et al. (1977) isolated a new paramyxovirus from R. f. ass¢milis and R. l. cooktownensts, and from mites of this species found on the latter host. Laelaps aella Domrow Laelaps aella Domrow, 1973, Proc. Linn. Soc. N.S.W., 98: 65. Material. Three 99, Pseudomys gracilicaudatus, 98 km NW Bundaberg (24°31’, 151°28’), Qd, 10.i1.1975; five 99, P. nanus (Gould), 7 km SE Nourlangie Camp (12°49’, 132°42'), N.T., 13.vi.1975; one 2, P. nanus, 14 km S Nourlangie Camp (12°54’, 132°38'), N.T., 18.vi.1975; nine 92, P. nanus, 346 km S Darwin (15°36’, 131°7’), N.T., 1.vi.1975; one 9, P. nanus, 18 km NE Kimberley Research Station (15°33’, 128°6), W.A., 28.v.1975; one 2, P. nanus, 246 km E Derby (17°7, 125°43'), W.A., 21.v.1975; four 99, P. nanus (two rats), 165 km E Derby (17°6, 125°10 ) , W.A., 15.v.1975. Notes. The only previous record of this species was from the Northern Territory. The considerable extension of range both eastward and westward now noted takes in all but the western extreme (New Norcia, W.A.) of the combined ranges of the two known hosts, if indeed they are specifically distinct (Ride, 1970). Their mites are indistinguishable. Laelaps rothschildi Hirst Laelaps rothschildz Hirst, 1914, Trans. zool. Soc. Lond., 20: 325. Material. Seven 292, Melomys cervinipes (four rats), 32 km S Cooktown (15°45’, 145°18'), Qd, 7-9.xi.1974; 18 99, M. cervinipes (four rats), 40-41 km SE Cairns Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 197 (17°15', 145°56’), Qd, 15-16.xi.1974; four 99, M. cervinipes, 19 km SE Atherton (17°25', 145°31'), Qd, 25.xi.1974; one 9, M. cervinipes, 61 km N Rockhampton (22°51, 150°40’), Qd, 5.1.1975; two 99, M. cervinipes, 98 km NW Bundaberg (24°32', 151°28'), Qd, 12.i1.1975; 21 99, M. lettoralds (six rats), Iron Range, 21-26 km S Portland Roads (12°44-48', 143°16-18’), Qd, 1-3.xi.1974; nine QQ, M. lettoralis (four rats) , Mount Simon, 22 km S Cooktown (15°39’, 145°13’), Qd, 8.xi.1974; three QQ, M. littoralis (two rats) , 37 km S Cooktown (15°48', 145°15’), Od, 7.xi.1974; two QQ, M. littoral’s, 25 km N Atherton (17°3’, 145°26') , Qd, 21.xi.1974; one 9, one pn, M. littoral’s, 11 km NE Atherton (17°12', 145°33’), Qd, 22.xi.1974; 12 99, M. littoralis (five rats), 17 km S Cairns (17°5’, 145°47°), Qd, 14.xi.1974; one 9, M. littoralts, 29 km SE Innisfail (17°46', 146°7'), Qd, 3.xii.1974; three 29, M. lttoralis (two rats), 58 km N Rockhampton (22°52’, 150°41’), Qd, 5.1.1975; three 99, M. littoralts, 58 km N Maryborough (25°6 , 152°32’), Qd, 14.1.1975; 13 99, one dn, one pn, M. lttoralds (three rats) , 9 km SE Dunwich, Stradbroke Island (27°32', 153°30)), Qd, 19-20.i1.1975; three 22, M. lttoralis, Leanyer Swamp, 15 km NE Darwin (12°22’, 130°56'), N.T., 11.vi.1975; two 92, Melomys sp., Kowanyama; six 9°, Melomys sp., 62 km NW Coen (13°27', 142°57'), Qd, 29.x.1974. Notes. This species is common on Melomys in New Guinea and coastal N.E. Australia (Domrow, 1973) , but was not previously recorded from the Northern Territory. Laelaps pammorphus Domrow Laelaps pammorphus Domrow, 1973, Proc. Linn. Soc. N.S.W., 98: 69. Material. Seventeen 99, two dd, Zyzomys argurus (Thomas) (six rats), Mount Simon, 22 km S Cooktown (15°39’, 145°13’), Qd, 7-9.xi.1974; eight 99, two dd, Z. argurus (five rats), Torola Pool, Fortescue River (21°18', 116°11), W.A., 8.v.1975; 29 29, three DG, Z. argurus (11 rats), 165 km E Derby (17°6-7', 125°10'), W.A., 15- 17.v.1975; six QQ, Z. woodward? (Thomas) (three rats), Canon Hill, 225 km E Darwin (12°23', 132°56’), N.T., 21.vi.1975. Notes. The only previous records of this species were from the Northern Territory. The considerable extension of range both eastward and westward now noted takes in all but the westernmost portion (the Pilbara, W.A.) of the combined ranges of the two known hosts (Ride, 1970). The strengthened setae J; and Z, originally noted on the dorsal shield of specimens from Z. woodward: are visible at x30 in spirit when suitably lit. Laelaps janalis, n. sp. (Figs 8-9) Types. Holotype 9 and four paratype 99, Pseudomys occidentalis Tate (two rats) , 17 km NE Bendering (32°21-22’, 118°28'), W.A., 1.iv. 1975; one paratype 2, same data as holotype, but 30.i11.1975. Female. Basis capituli longer than wide, with setae c short, about one-quarter as long as interspace, falling short of sides of basis; deutosternum with six denticles mostly in single file, but first and last denticle at times multiple. Hypostome with setae h3>h,>h; h, about one-fifth longer than interspace; h, short, = c; h; almost twice as long as interspace, well exceeding sides of basis. Labial cornicles well formed. Labrum spiculate, hastate. Epistome soft and diaphanous, apparently with two small lobes in median indentation. Palpal setation (trochanter-tibia) holotrichous, i.e. 2.5.6.14 (including two dorsodistal tibial rods) ; seta v, on trochanter elongate, slightly flared; seta al, on genu slightly spatulate; tarsus with one of three v elongate; claw bifid, tines with extended, hyaline, minutely barbed ‘ edges. Chelicerae with basal segment subequal in diameter to, but only half as long as, shaft of distal segment; digits occupying one-fifth of total length. Fixed digit with incurved tip and one distal tooth, Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 198 SOME DERMANYSSID MITES (ACARI) pp P Sf. f ? |} —— p ~~}. Figs 8-9. Laelaps janalzs. Idiosoma in dorsal and ventral views, @. between which is set short, stiff pilus dentilis about as long as diameter of digit at that level; with subbasal seta dorsally, but associated pores not detected. Movable digit with incurved tip and two external teeth, between which is accepted armature of fixed digit. Corona comprised of about ten subequal ciliations. Idiosoma 1,230-1,375 um long, 900-1,045 ym wide (not gravid) ; 1,440 um long, 1,080 ym wide (carrying fully developed larva) . Dorsal shield well sclerotized, surface marked by paired muscle insertions and with obvious reticulation; with usual sinuous vertical and humeral margins, with sides then very slightly diverging to two-thirds length, and finally converging rather more sharply to truncate posterior margin; podonotum with normal 22 pairs of setae, mostly short (J:-6, 21-6, 51-6, 12-5); opisthonotum with normal 17 pairs of setae, all short except Zs (J:-5, Zi-s, S:-s, PX2-3) ; pores in 22 pairs, those in front of S,.; overlain by extensive patch of transparent cuticle. Dorsal cuticle sclerotized except for narrow marginal strip, with about 12 pairs of setae of increasing length posteriorly. Tritosternal base unarmed; laciniae lightly ciliated, reaching forward to insertions of labial cornicles. Sternal shield strongly sclerotized, especially a broad band on anterior and lateral margins, and with heavy cornua between coxae I-II; surface without any obvious reticulation except laterally; anterior margin roundly convex; posterior margin shallowly concave, without any median extension; shield with three pairs of short, slender setae and two pairs of pores (each provided with Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 199 canaliculus). Metasternal shields small, each bearing small seta (provided with canaliculus), but pore free in adjacent cuticle. Genitoventral shield strongly sclerotized, with cuticle encroaching on lateral margins; surface marked by paired muscle insertions, but without obvious reticulation; shield expanded behind coxae IV, with sides subparallel and posterior margin shallowly concave; with pair of short genital setae and three pairs of usurped ventral setae (first pair short, other two pairs long), but pores free in adjacent cuticle; operculum broadly arched, sclerotized internally and supported by strong genital apodemes. Anal shield well sclerotized except for elongate clear central patch bearing anus and setae; lateral angles each - with muscle insertion and marginal pore; shield slightly longer than wide, with both antero- and posterolateral margins slightly concave; adanal setae set just behind anus, falling well short of long postanal seta, and latter far exceeding cribrum. Metapodal shields and two pairs of shieldlets between them and genitoventral shield all largely encroached on by cuticle. Peritremes of medium length, extending forward only to level of anterior margins of coxae II, borne on peritrematal shields that run forward to fuse with dorsal shield vertically but are free of broadly crescentic exopodal shields IV behind. Ventral cuticle with about 30 pairs of setae of increasing length posteriorly, including one long pair immediately behind genitoventral shield. Legs with setation called for by Evans and Till (1965) for dermanyssids in general, and for L. echidninus in particular (i.e. holotrichous except for one additional pl seta on genu IV, 2-5/1-2). Coxa III with pu a heavy spine. Trochanters I-II with al, and III-IV with al and d strengthened, but still sharply pointed. Femora I-II with pd, lengthened, reaching distal margin of tibia in former, but barely as long as basal diameter of segment in latter; II with av slightly, and III with wv heavily, spinose. Genu I with pd; lengthened, a little longer than ad, on femur I. Tarsi II-IV with ad, and pd, minute and other setae, especially ventrodistally, strengthened (in particular, al,, av, and pl, on II and al,_, on III) ; II without ad; unduly lengthened as in L. albycta Domrow, 1965. Larva. Details not clear within 2, but podonotum holotrichous, with 10 pairs of setae ranging from 90 (j,) to 290 um (j,) in length; opisthosoma also with several pairs of elongate setae. Notes. This fine new species will not go beyond the first couplet in Domrow’s (1965, 1973) keys and diagnoses. In showing a relatively full complement of setae on the dorsal shield and peritremes of at least medium length, it fits with the nuttallz, spatanges and hapaloti groups; but the minute genital and first pair of usurped ventral setae on the genitoventral shield indicate the f¢nlaysonz group. A review of the Australian Laelaps spp. awaits study of a collection from the Kimberley region of Western Australia (this includes further new species, but carnot be treated here by reason of the conditions of loan) . The specific name is a Latin adjective, janal-zs, -e, of Janus, the Roman god of the year, who looked both to the front and back. Laelaps bycalza, n. sp. (Figs 10-17) Types. Holotype 2, allotype 3, two paratype 92, one morphotype dn and one morphotype pn, Pseudomys albocinereus (Gould) (two rats), 22 km NE Jurien (30°8', 115°9’), W.A., 13.iv.1975. . Female. Capitulum as in L.albycta except as follows. Setae c shorter, about one- quarter as long as interspace, falling short of sides of basis. Setae h, slightly longer; hs longer, about three-quarters as long as interspace. Epistome with four small, weak median lobes. Basal segment of chelicerae one-tenth shorter than shaft of distal segment. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 200 SOME DERMANYSSID MITES (ACARI) 10 3 Figs 10-13. Laelaps bycalta. 10-11. Idiosoma in dorsal! and ventral views, 2. 12-13. Idiosoma in dorsal and ventral views, 0. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 201 Idiosoma 1,100 um long, 905 um wide (older specimen bearing larva) ; 990 um long, 825 wm wide (recently moulted, non-gravid specimen). Dorsal shield moderately well sclerotized, surface marked by paired muscle insertions but without reticulation; outline essentially as in L. janalis, but tapering more sharply in posterior third, more sharply truncate posteriorly and of rather broader proportions; podonotum with 20 pairs of setae, mostly short (first two, somewhat isolated pairs on dorsal cuticle could well be 73_,, thereby making up normal 22 pairs) ; opisthonotum with 15 pairs of setae, all short except Z; (px2-3 lacking) ; pores as in L. janalis, but patches of transparent cuticle in front of S,.; smaller. Dorsal cuticle sclerotized except for marginal strip (evident in recently moulted specimen only as slight granulation of cuticle), with about 12 pairs of setae of slightly increasing length posteriorly (excluding two isolated anterior pairs noted above) . Venter as in L. janalis except as follows. Sternal shield with posterior margin almost straight and all setae rather longer, st; well exceeding posterolateral angles. Genitoventral shield with outline almost straight between genital setae and first pair of usurped ventral setae, then rounding sharply to ever so slightly concave posterior margin; difference between two types of setae on shield more marked. Anal shield with clear central patch not extending to cribrum; anterior margin strongly arched ; anal setae smaller, especially adanals. Peritremes abbreviated, situated entirely above coxae III. Ventral cuticle marginally with about seven pairs of setae of increasing length posteriorly and two marked pairs immediately behind genitoventral shield. Legs with setation called for by Evans and Till (1965) for dermanyssids in general, and for L. echidninus in particular (i.e. holotrichous except for one additional pl seta on genu IV, 2-5/1-2). Details as in L. janalzs except on tarsi: II-III (but not IV) with setae, especially ventrodistally, strengthened; II as in L. albycza, i.e. al, and pl, strengthened (but still pointed), ad; and (to lesser extent) av; lengthened, and one seta on basitarsus (ad,) strengthened; III with al,_,, av; and (on basitarsus) al, strengthened. Male. Capitulum as in 2 except as follows. Some deutosternal denticles multiple. Hypostomatal setae subequal, h, about two-thirds as long as interspace. Labial cornicles softer, flared at tips. Chelicerae with fixed digit soft, gently tapering and edentate, 55 um long, 8 um wide basally. Spermatodactyl 110 wm long, occupying five-sixteenths of total cheliceral length; lightly upcurved, with edentate, but firmly pointed, remnant of movable digit just beyond half length. Neither chelicera protruded far and corona not detected. Idiosoma 890 um long, 705 wm wide (slightly ruptured). Dorsum as in 9 except as follows. Dorsal shield less angulate in posterior third; podonotum with 21 pairs of setae, i.e. taking in 73, but presumptive r, (lacking on one side) still free in cuticle; also irregular in lacking one z, and one S,, and in doubling of one Z,. Strip of sclerotized cuticle narrow, irregular and largely incomplete posteriorly. Venter as in 2 except as follows. Holoventral shield reticulate except for elongate patch extending from level of metasternal setae to disc of genitoventral portion; setae st,_3 and metasternal setae far longer; genitoventral portion with genital setae and five pairs of usurped ventral setae, difference between two types of setae less marked. Metapodal shields simpler. Leg setation as in 9 except as follows. Coxa IV with v much longer. Some other setae stronger, resembling pu on coxa II]: femora I four v, II pu,, IV v; genua I, Il pv, Lav and pv, IV pl,; tibiae I, III-IV pu, Il av and pv. Deutonymph (enclosing 6). Capitulum as in 6 except that chelicerae resemble those of 2. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 202 SOME DERMANYSSID MITES (ACARI) Figs 14-17. Laelaps bycalia. 14-15. Idiosoma in dorsal and ventral views, dn. 16-17. Idiosoma in dorsal and ventral views, pn. Idiosoma 790 um long, 620 ym wide. Dorsum as in Q, but shield less angulate in posterior third, with longer setae (r; free in cuticle on one side) , showing only traces of lateral incisions between podonotal and opisthonotal halves, and not invested by strip of sclerotized cuticle. Venter with same setation as 6, but sternogenital shield discrete and peritrematal shields less developed. Leg setation anticipating that of 6, including genu IV. Protonymph. Capitulum holotrichous, anticipating that of 9. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 203 Idiosoma 670 um long, calculated to be 550 ~m wide. Podonotal shield with three shallow lobes posteriorly; podonotum holotrichous, with 16 pairs of setae (eleven on shield, five on cuticle). Pygidial shield transverse, slightly concave anteriorly but with slight median convexity; opisthonotum normally holotrichous, with 14 pairs of setae (eight on shield, six on cuticle — superficial count would give thirteen pairs, but closer examination reveals S, lacking on one side of shield and S; on other). Sternal shield elongate, with posterior margin distinctly triangulate; with usual three pairs of setae and two pairs of pores. Genital complex represented only by pair of distinct pores. Ventral cuticle with four pairs of setae. Peritrematal shields in three fragments. Leg setation holotrichous except for one additional p/ seta on genu IV (1-4/0-1), predicting that of 9 except for coxae: III with pu hardly strengthened, IV with v elongate. Larva. Details not clear within 9, but podonotum holotrichous, with 10 pairs of setae ranging from 18 (s.) and 25 (7,) to 135 ym (7,) in length. Opisthosoma with a few pairs of setae resembling s.. Notes. This new species keys out near L. ftnlaysont Womersley, 1937, see Domrow (1963, 1965), but the outlines of both the dorsal and genitoventral shields in the female show clear differences. Further, although both species show 35 pairs of setae on the dorsal shield, there also appear to be fundamental differences in the J and px series on the opisthonotal portion; these await further study, see notes on L. janalis above. The specific name of this, and the other new species below, are both anagrams of calabyz, and are to be treated as nouns (nominative singular) in apposition to Laelaps. Laelaps cybiala Domrow Laelaps cybiala Domrow, 1963, Proc. Linn. Soc. N.S.W., 88: 206. Material. Two 99, Mastacomys fuscus Thomas, Wragge Creek, Kosciusko National Park (36°23', 148°28’), N.S.W., 10.11.1975. Notes. This material confirms the original record. Laelaps lybacza, n. sp. (Figs 18-19) Types. Holotype 2, Pseudomys praeconis Thomas, Bernier Island (24°55', 113°8’), WiAve 22 1ve 1 9175). ; Female. Capitulum as in L. albycia except as follows. Setae c much shorter, about one-eighth as long as interspace, falling well short of sides of basis. Setae A, not fully clear, but shorter than interspace; h; longer, about three-fifths as long as interspace. Epistome not clear. Basal segment of chelicerae one-fifth shorter than shaft of distal segment. Idiosoma 1,035 um long, 845 um wide. Dorsal shield moderately well sclerotized, surface marked by paired muscle insertions but without reticulation except for two or three weak humeral lines; outline intermediate between those of L. janalzs and L. bycalta; podonotum with 19 pairs of setae, mostly short (z; lacking, first of two, somewhat isolated pairs on dorsal cuticle could well be 73.4) ; opisthonotum with 14 pairs of setae, all short except Z; (S; lacking, J and px series requiring further study) ; pores as in L. janalis, but patches of transparent cuticle in front of S,-5 slightly smaller. Dorsal cuticle sclerotized except for broad marginal strip, with about 17 pairs of setae of slightly increasing length posteriorly (excluding two isolated anterior pairs noted above). Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 204 SOME DERMANYSSID MITES (ACARI) Figs 18-19. Laelaps lybacza. Idiosoma in dorsal and ventral views, @. Venter as in L. bycalia except as follows. Sternal shield with posterior margin slightly concave and setae st, short compared to st,_3. Genitoventral shield rounded laterally and more concave posteriorly. Adanal setae stronger, postanal weaker. Legs with setation called for by Evans and Till (1965) for dermanyssids in general, and for L. echzdnznus in particular (i.e. holotrichous except for one additional p/ seta on genu IV, 2-5/1-2). Details asin L. bycalza. Notes. This new species keys out near L. finlaysonz, see Domrow (1963, 1965), but differs therefrom in lacking setae z; and S$; on the dorsal shield, and in its broader genitoventral shield. The J and px series on the dorsal shield show certain funda- mental similarities, but await further study, see notes on L. janalis and L. bycalea above. Eulaelaps stabularts (Koch) Gamasus stabularts Koch, 1839, Deutschlands Crustaceen, Myriapoden und Arachniden. Regensburg: Herrich-Schaffer. Heft 27: No. 1. Material. Two 29, Felis catus Linnaeus, Hamilton, Vic., xi.1975, A.M. Freemantle. Notes. The only other record of this apparently recently introduced Holarctic and Oriental species in Australia is from Mus musculus Linnaeus in Tasmania (Domrow, 1973). Echinonyssus butantanensis (da Fonseca) Ichoronyssus butantanensis da Fonseca, 1932, Mem. Inst. Butantan, 7: 135. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 R. DOMROW 205 Material. One @, Rattus rattus, 26 km NE Perth (31°13, 116°9’), W.A., 4.iv.1975. Notes. Previous Australian records of this introduced species were all from the east: Womersley (1956, as Hirstzonyssus arcuatus (Koch); Glenfield is near Sydney, N.S.W., not in Qd) and Domrow (1961, 1963, as H. musculi (Johnston) ). The specific name now used is after Evans and Till (1966) and Herrin (1974). Figs 20-21. Trichosurolaelaps crasstpes. \diosoma in dorsal and ventral views, pn: Trichosurolaelaps crasstpes Womersley (Figs 20-21) Trichosurolaelaps crasstpes Womersley, 1956, Linn. Soc. J., Zool., 42: 564. Materzal. Six 29, Trichosurus vulpecula (Kerr), Belbora, Kowanyama, iv.1969, R. Domrow and E. T. Bulfin; three 292, two dd, T. vulpecula, Kowanyama; three @, T. vulpecula, D’Aguilar, Qd, 1.iv.1957, R. Domrow; one G, T. vulpecula, Grovely, Qd, 24.11.1965, E. H. Derrick; eight 92, one d, T. vulpecula, Brookfield, Qd, 30.i11.1973, G. Wolf; six 99, T. vulpecula, Woodridge, Qd, 29.x.1963, R. Domrow and I. D. Fanning; three 92, T. vulpecula, Bonalbo, N.S.W., 2.vi.1961, K. Keith and D. L. McIntosh; two dd, one dn, one pn, T. vulpecula, Taronga Park Zoo, Sydney, N.S.W., 18.viii.1967, M. D. Murray; five 29, five Jd, four dn, four pn (sample only), T. vulpecula, Sunbury, Vic., v.1977, J. H. Arundel; one 2, three dG, T. vulpecula, North Midlands, Tas., 2.vii.1962, R. H. Green; one 9, one G, T. vulpecula, Kelso, Tas., 16.11.1961, B. C. Mollison; eight 99, two dd, T. vulpecula, Maydena, Tas., 10.v.1961, B. C. Mollison. Deutonymph. Details of capitulum and legs as in adult, but armature of legs weaker, betraying its setal origin (e.g. in prefemale, more than in premale, seta av, on genu- tibia I is spinose basally, but then strongly notched and setiform distally) . Idiosoma 450-460 um long, 285-290 um wide (prefemale) ; 445-470 um long, 295-310 um wide (premale). Dorsal shield not incised laterally between podonotal and opisthonotal portions; setation as in 3, but setae on margin of shield (behind level of coxae II) and those on cuticle distinctly bladed. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 206 SOME DERMANYSSID MITES (ACARI) Sternogenital shield with four pairs of setae and three pairs of pores, gently tapering behind setae st, to terminate roundly between genital setae (genital pores not detected). Anal shield as in 2. Setae of ventral cuticle bladed, in eight to ten pairs. Peritremes much abbreviated both in prefemale and in premale, barely as long as diameter of stigmata (characteristic adult form clear in one enclosed @) . Protonymph. Palpal trochanter-genu holotrichous. Idiosoma 310-340 um long, 190-220 um wide (in front of peritremes) . Podonotal shield trilobed posteriorly, median lobe the strongest; podonotum holotrichous, with 16 pairs of setae (11 on shield, five on cuticle — j,_; and z, of differing lengths; 74.., z4-s and s, minute; ss; and five pairs on cuticle long and bladed). Pygidial shield semicircular, but with median convexity on anterior margin; opisthonotum bideficient, with 12 pairs of setae (six on shield, six on cuticle — J,_; minute, as is submarginal Z, between marginal S,_; of medium length; Z; tong but simple; six pairs on cuticle long and bladed) .* Sternal shield with usual three pairs of setae (of increasing length posteriorly; st, slightly, and st; clearly, bladed) and two pairs of pores. Metasternal and genital complexes not detected. Anal shield predicting that of 9. Ventral cuticle with three pairs of bladed setae in front of, and one pair of stout setae flanking, anal shield. Stigmata strong, protuberant, without peritremes. Leg setation holotrichous except that tibia IV is unideficient posterolaterally (1-3/2-0) . Armature already predicting that of adult. Notes. These records document Domrow’s (1972) bald statement that this species is common on this host in eastern Australia. Kowanyama is the northernmost record. Troughton (1965: 105) noted that small [unidentified] mites caused annoyance and irritation in Schoznobates volans (Kerr), especially when sickly. Species of Trichosurolaelaps may now be linked with tissue damage in their hosts, at least in zoo conditions. The series of T. crass¢pes from Taronga Park Zoo stemmed from areas of alopecia and keratinization around the head, elbows, hocks and perineum, with evidence of intense itching; that from Sunbury from the back of a mature female possum that, having been held captive for six months, had been biting at the area for two to three weeks, the mites apparently being quite irritating. The series of T. strzatus below from Dandenong was from a skin scraping; that from Werribee from a heavily infested possum with a large area of hair loss consistent with trauma induced by irritation. Trichosurolaelaps striatus Domrow Trichosurolaelaps striatus Domrow, 1958, Proc. Linn. Soc. N.S.W., 82: 356. Material. Many specimens, Pseudocheirus peregrinus (Boddaert) , Mosman, N.S.W., 26.v.1966, A. L. Dyce; two 99, P. peregrinus, Dartmouth, Vic., 22.xi.1973, I. Beveridge; three 99, one 6, P. peregrinus, Dandenong, near Melbourne, Vic., vii.1977, N. J. Barton; many specimens, P. peregrinus, Werribee, Vic., 1x.1977, J. H. Arundel. Notes. Domrow (1961) extended the range of this species from S.E. Queensland to Tasmania, but without intermediate records. See also notes on preceding species. *Setae J, and Z, are assigned these signatures because the former is set just inside a wideset pair of pores (cf. holotrichous condition in Figs 1 and 16), and the latter because its position allows only this interpretation. Domrow (1972), writing before immatures were known in this genus, but knowing that J3_4 and Z;., all occur in the holotrichous condition, arbitrarily considered the posteriormost seta possible to be the absent one in the case of deficiencies in the adults. When further immatures are known, the situation can again be analysed. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 . R. DOMROW 207 Ornithonyssus bacoti (Hirst) Letognathus bacot2 Hirst, 1913, Bull. entomol. Res., 4: 122. Material. Two pn, Rattus rattus, 26 km NE Perth (31°13', 116°9’), W.A., 4.iv.1975. Notes. This species, the tropical rat mite and a minor pest of man, is widespread in Australia (Hirst, 1914; Domrow, 1963, 1973). Closer examination quickly showed that these two nymphs did not belong with the female of E. butantanensis above that was collected on the same rat. 22 23 7 \ / a Ng ) Figs 22-23. Ornithonyssus syluiarum. 22. Six variants of tapered posterior portion of dorsal shield, 9 (four pairs of longer, and one pair of shorter, setae are normal complement; 12 pairs more anteriorly). 23. Five variants of sternal shield, 9 (second from bottom most typical) . Ornithonyssus syluiarum (Canestrini and Fanzago) (Figs 22-23) Dermanyssus syluarum Canestrini and Fanzago, 1877, Attz Ist. Veneto, 5: 124. Material. Seven 29, one pn, nestling Hirundo neoxena Gould, Campania, Tas., Sr LOVA Ie Park. Notes. These specimens, from a new host, underline the widespread occurrence of this pest of poultry on native birds in temperate southern Australia (Domrow, 1973). A customary key character for this species is the disassociation of setae st; from the sternal shield, but the condition varies considerably from specimen to specimen (as does the setation of the posterior portion of the dorsal shield) ; see also Allred (1970). Halarachne miroungae Ferris Halarachne miroungae Ferris, 1925, Parasctology, 17: 166. Matertal. Two 1, Mirounga leonina (Linnaeus) , Macquarie Island, Southern Ocean, summer of 1976, I. Morgan. Notes. The previous nearest record of this species to Australia probably originated from Kerguelen, some 5,000 miles to the west (Domrow, 1962b, 1974) . Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 208 SOME DERMANYSSID MITES (ACARI) References ALLRED, D. M., 1970. — Dermanyssid mites of New Guinea. J. med. Entomol., 7: 242-246. CAMPBELL, R. W., Car ey, J. G., DoHERTy, R. L., Domrow, R., FitippicH, C., Gorman, B. M., and Karasatsos, N., 1977. — Mossman virus, a paramyxovirus of rodents isolated in Queensland. Search, 8: 435-436. Domrow, R., 1958. — New and little known Australasian Laelaptidae (Acarina). Proc. Linn. Soc. N.S.W., 82: 352-366. ——, 1961. — New and little-known Laelaptidae, Trombiculidae and Listrophoridae (Acarina) from Australasian mammals. Proc. Linn. Soc. N.S.W., 86: 60-95. —, 1962a. — Mammals of Innisfail. II. Their mite parasites. Aust. J. Zool., 10: 268-306. ——, 1962b. — Halarachne miroungae Ferris redescribed (Acarina: Laelaptidae). Pac. Insects, 4: 859- 863. —— , 1963. — New records and species of Austromalayan laelapid mites. Proc. Linn. Soc. N.S.W., 88: 199-220. —, 1965. — The genus Laelaps in Australia (Acarina: Laelapidae). J. entomol. Soc. Queensl., 4: 18- 23. ——, 1966. — Some laelapid mites of syndactylous marsupials. Proc. Linn. Soc. N.S.W., 90: 164-175. ——, 1967. — Mite parasites of small mammals from scrub typhus foci in Australia. Aust. J. Zool., 15: 759-798. ——, 1972. — The crasstpes species-group, genus Trzchosurolaelaps Womersley (Acari: Dermanyssidae) . J. Aust. entomol. Soc., 11: 295-305. ——, 1973. — New records and species of Laelaps and allied genera from Australasia (Acari: Dermanyssidae). Proc. Linn. Soc. N.S.W., 98: 62-85. ——, 1974. — Notes on halarachnine larval morphology and a new species of Pneumonyssus Banks (Acari: Dermanyssidae) . J. Aust. entomol. Soc., 13: 17-26. ——, 1977. — New records and species of Laelaps and allied genera from Australasia (Acari: Dermanyssidae). Part 2. Proc. Linn. Soc. N.S.W., 101: 185-217. Evans, G. O., 1969. — Observations on the ontogenetic development of the chaetotaxy of the tarsi of legs II-IV in the Mesostigmata (Acari). Proc. II int. Congr. Acarol., 1967: 195-200. ——, and Titi, W. M., 1965. — Studies on the British Dermanyssidae (Acari: Mesostigmata). Part I. External morphology. Bull. Br. Mus. nat. Hist., Zool., 13: 247-294. ——, 1966. — Studies on the British Dermanyssidae (Acari: Mesostigmata). Part II. Classification. Bull. Br. Mus. nat. Hist., Zool., 14: 107-370. HERRIN, C. S., 1974. — The taxonomic status of Hirstionyssus butantanenses (Fonseca, 1932) (Acari: Mesostigmata) . J. med. Entomol., 11: 341-346. Hirst, S., 1916. — On the occurrence of the tropical fowl mite (Lzponyssus bursa, Berlese) in Australia, and a new instance of its attacking man. Ann. Mag. nat. Hist., (8) 18: 243-244. MarsHALL, A. G., 1976. — Host-specificity amongst arthropods ectoparasitic upon mammals and birds in the New Hebrides. Ecol. Entomol., 1: 189-199. MITCHELL, C. J., 1964. — Ectoparasitic and commensal arthropods occurring on the rats of Manoa Valley, Oahu (Acarina, Anoplura, and Siphonaptera) . Proc. Hawazz. entomol. Soc., 18: 413-415. Rive, W. D. L., 1970. — A Guide to the native Mammals of Australia. Melbourne: Oxford University Press. ROBINSON, J. F., RoBinson, A. C., Watts, C. H. S., and BaversTock, P. R., 1978. — Notes on rodents and marsupials and their ectoparasites collected in Australia in 1974/75. Trans. R. Soc. South Aust., 102: 59-70. Tay or, J. M., and Horner, B. E., 1973. — Results of the Archbold Expeditions. No. 98. Systematics of native Australian Rattus (Rodentia, Muridae). Bull. Am. Mus. nat. Hist., 150: 1-130. TENORIO, J. M., and Rapovsky, F. J., 1974. — The genus Mesolaelaps (Laelapidae: Mesolaelapinae, n. subfam.) with descriptions of two new species from New Guinea. J. med. Entomol., 11: 211-222. TROUGHTON, E., 1965. — Furred Animals of Australia. Sydney: Angus & Robertson. Eighth Ed. WomersLey, H., 1937. — Studies in Australian Acarina Laelaptidae. I. New records and species of Laelaps and allied genera. Parasitology, 29: 530-538. ——, 1956. — On some new Acarina-Mesostigmata from Australia, New Zealand and New Guinea. Linn. Soc. J., Zool., 42: 505-599. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 Feeding Habits and Structure of the Gut of the Australian Freshwater Prawn Paratya australiensis Kemp (Crustacea, Caridea, Atyidae) PREM GEMMELL GEMMELL, P. Feeding habits and structure of the gut of the Australian freshwater prawn Paratya australiensts Kemp (Crustacea, Caridea, Atyidae). Proc. Linn. Soc. N.S.W. 103 (4). (1978) 1979: 209-216. A study of the feeding habits and histological details of the alimentary canal of Paratya australiensis is presented. Functional morphology and histology of the various regions of the gut are described and discussed in relation to food and feeding habits. Prem Gemmell, School of Anatomy, University of New South Wales, Kensington, Australia 2033; revised manuscript received 27 March 1978, accepted 20 September 1978. INTRODUCTION Alimentary adaptations in decapod crustaceans have attracted the attention of workers since the beginning of the century. Parker (1876), Calman (1909), Yonge (1924) , and Patwardhan (1934, 1935) were pioneers in the field. Recently George et al. (1955), Fryer (1960), Schaefer (1970) and Powell (1974) have contributed considerably to the study of decapod alimentary organization. Paratya australiensis (Kemp) is a common freshwater small prawn that favours vegetated areas. It occurs in a wide variety of permanent inland waters (coastal streams, rivers, lakes, farm dams and ponds). Williams (1977) described its occurrence in south-eastern South Australia, south-western New South Wales extending northwards into Queensland. The present investigation is the first study of the alimentary adaptations of this species in relation to its feeding habits. MATERIALS AND METHODS Specimens of Paratya australiensis were collected monthly from Manly Dam, N.S.W., during the period October 1972 to February 1973. The specimens were brought to the laboratory alive and were kept at a constant temperature of 18°C for further observations. For histological studies specimens were fixed in alcoholic Bouin’s fluid for 24 hours. The antennules, antenna, thoracic and abdominal appendages were cut right to the base and the carapace was completely removed to allow easy penetration of the fixative. The fixed material was subsequently washed, dehydrated, embedded in paraffin (M.P. 56°C) and sectioned at 6 um. The sections were stained in Delafield haematoxylin and eosin and mounted in D.P.X. mounting medium. OBERVATIONS FEEDING HABITS Paratya australiensis is a browser and filter feeder. During browsing the food is collected by chelipeds. The food which consists of fine particles and insects is scraped off the substratum by the strong toothed spines. As the fingers of the chela approach the substratum, they open widely; on contact with the substratum the terminal spines Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 210 AUSTRALIAN FRESHWATER PRAWN and setae become splayed out. As the chela closes and begins to be withdrawn its setae come together; their elasticity ensures that the extremities are held closely to the substratum and a considerable area is scraped. The scraped material is held by the opposed sets of setae. The serrated edges of the toothed spines of chela are responsible for scraping the substratum and loosening the detritus. Larger particles are picked up by the combed, and the finer ones by the brush setae. The loaded spines then pass into the mouth parts where the grid setae of the first maxilliped strip off the food. The chelipeds are also used for breaking down the larger particles of food. During filter feeding, Paratya holds itself in a slanting position near the surface of the tank. Strong water currents are produced by the metachronal beatings of the endites of the first maxilla and scaphognathite of the second maxilla, these appendages are fringed with long dorsally directed setae which pass the food in the midventral line towards the mouth. The direction of food currents during this action is firstly towards the animal from the sides and secondly, from behind forwards along the mid-ventral line. Small particles of food suspended in water are taken up by the toothed and plumose setae on the inner side of the endites of the first and second maxillipeds. The stout spinous setae of the first maxilla help in scraping off the food from the setae of the more posterior appendages. The food is passed into the buccal cavity via the mandibles. Fig. 1. T.S. through the oesophagus of Paratya australiensis showing the disposition of the oesophageal folds. C = chitin, C.M.F. = circular muscle fibres, D.F. = dorsal fold, EP. = epithelium, L.M.F. = long muscle fibres, M = mouth, MAN. = mandible, OES. = oesophagus, SM. = submucosa. Magnification : approx. x90. Fig. 2. T.S. through mouth, oesophagus and the cardiac proventriculus showing the cardio-oesophageal junction. C = chitin, C.OES.O = cardio-oesophageal orifice, C.PROV. = cardiac proventriculus, D.F. = dorsal fold, L.F. = lateral fold, P. PROV. = pyloric proventriculus, S = seta. Magnification: approx. x90. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 P. GEMMELL 211 STRUCTURE OF THE GUT The alimentary canal of Paratya australzenszs consists of a foregut, midgut and hindgut. FOREGUT The foregut includes the mouth, oesophagus and the tventriculus. The mouth (M’) is ventral between the two mandibles. In front the labrum overlaps the mouth and the incisor processes of the mandibles. The oesophagus (Fig. 1) is a narrow tube ascending from the mouth and opening into the cardiac proventriculus. The cardiac end of the oesophagus is dilated and thrown into three folds — one dorsal and two lateral (D.F. and L.F.). The wall of the oesophagus is lined internally by chitin, columnar epithelium, thick submucosa of loose connective tissue, circular and longitudinal muscle layers and an external layer of connective tissue. The proventriculus (Fig. 2) is divided into an anterior large cardiac proventriculus (C.PROV.) and a posterior small pyloric proventriculus (P.PROV.). The two are separated by a prominent depression. The cardiac proventriculus is a dilated pouch in which food can be accumulated. Regurgitation of food from the cardiac chamber into the oesophagus is prevented by the oesophageal folds (D.F. and L.F.) at the termination of the oesophagus. The cuticular lining (C.) of the cardiac proventriculus constitutes a setose gastric mill (G.M.) The cardiac proventriculus in Paratya shows a median vertical bar (M.V.B.) extending from the anterior to nearly the posterior end of the chamber. Two pairs of short horizontal bars (H.B.) attached to its sides are covered with chitin (C.). The lateral bars show serrated edges on both sides; probably to increase the triturating surface. The chitinous lining (C.) of the cardiac proventriculus is produced into long fine setae (S.) which are inwardly and downwardly directed (Fig. 2). The chitinous lining of the pyloric proventriculus is produced into a number of setae (S.) forming the filter apparatus (Figs 3 and 4). From the base of the pyloric proventriculus arises a median ventral piece (M.V.P.) that extends up to the anterior end dividing the pyloric proventriculus into two narrow lateral grooves (L.G.) ; one on either side of the median piece. The chitinous lining of the pyloric proventriculus is produced into a number of spines (S.) all along its lateral edges. The spines are inwardly and upwardly directed, becoming progressively smaller from the base to the apex. The lateral edges of the median ventral piece (M.V.P.) are similarly produced into small groups of spines at regular intervals. The spines are large at the base, becoming smaller towards the apex. The larger spines overlap the smaller ones in front of them producing a many-pocketed sieve-like structure. Between the two sets of spines, that is the ones along the lateral lining of the pyloric chamber and those on the median ventral piece is a narrow space — the lateral groove (L.G.) which ultimately releases fine food particles into the midgut (M.G.) (Fig. 5). The coarse particles are retained for further action. This observation is supported by the presence of large food particles in the cardiac and fine in the pyloric proventriculus. The pyloric filter apparatus of Paratya is continued into the midgut in the form of chitinous pyloric sheath (P.S.). The sheath collects coarser food particles from the filter apparatus and prevents it from mixing with fine food particles. The pyloric sheath (P.S.) in Paratya consists of only two semicircular chitinous folds lying opposite each other lateromedially and slightly overlapping at the sides. The epithelium lining the proventriculus is tall columnar, supported by a thin basement membrane (B.M.) ; a submucosa of loose connective tissue, circular muscle fibres and a peripheral layer of loose connective tissue. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 212 AUSTRALIAN FRESHWATER PRAWN > RETIN Figs 3 & 4. T.S. through the proventriculus showing details of the gastric mill, filter apparatus and the disposition of the digestive glands. B.M. = basement membrane, C. = chitin, C.M.F. = circular muscle fibres, C. PROV. = cardiac proventriculus, C.P.O. = cardiopyloric orifice, EP. = epithelium, F.A. = filter apparatus, G.M. = gastric mill, H.B. = horizontal bar, INT. C. = intestinal caeca, L.G. = lateral groove, M.V.B. = median vertical bar, M.V.P. = median ventral piece, S = spine, SM. = submucosa. Magnification: approx. x110. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 P. GEMMELL 213 MIDGUT The midgut (Figs 5 and 6) is a short tube. Two long diverticulae — the intestinal caecae (INI.C.) arise from its anterior end and extend on either side of the proventriculus. The midgut is lined by columnar epithelium supported by a thin layer of loose connective tissue, an inner layer of circular muscle fibres and an outer layer of longitudinal muscle fibres followed by a thin investing sheath. HINDGUT The hindgut is short and narrow. The inner surface is thrown into a number of broad folds (Fig. 7). The wall consists of the following layers — an inner layer of columnar epithelium covered by chitin, submucosa of loose connective tissue, inner layer of circular (C.M.F.), outer layer of longitudinal muscle fibres (L.M.F.) anda thin limiting membrane. DIGESTIVE GLANDS The digestive glands are complex greenish-yellow tubular structures occupying most of the cephalothorax. The lobules of the glands are arranged around a lobular duct. These ducts join together to form the main duct that opens into the midgut. The lumen is lined by tall columnar secretory cells that are highly vacuolated, the resting cells showing a number of granules in the cytoplasm. The epithelium is supported by connective tissue and smooth muscles which encourage the discharge from the secretory units. Fig. 5. T.S. through the proventriculus and the mesenteron showing the opening of the filter apparatus into the mesenteron. B.M. = basement membrane, DG. = digestive glands, EP. = epithelium, INT. C. = intestinal caeca, MES. = mesenteron. Magnification: approx. x110. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 214 AUSTRALIAN FRESHWATER PRAWN Fig. 6. T.S. through the mesenteron showing the position of the pyloric sheath. C.M.F. = circular muscle fibres, P.S. = pyloric sheath. Magnification: approx. x110. Fig. 7. T.S. through the proctodaeum. C.M.F. = circular muscle fibres, EP. = epithelium, L.M.F. = longitudinal muscle fibres, SM. = submucosa. Magnification: approx. x250. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 P. GEMMELL 215 DISCUSSION The maxillae in Paratya australiensis are adapted for producing water currents. The endites of the first maxilla and the scaphognathite of the second maxilla along with the maxillipeds produce strong water currents. Smaller particles of food are picked up by the thickly arranged setae on these appendages and passed to the mouth. The chelipeds are used for picking up larger particles of food, breaking it into bits and their final transference to the mouth. The chelipeds are also responsible for scraping, collecting the detritus and its transfer to the mouth. The present investigations are in accord witth the finds of Fryer (1960) on Carzdina africana and C. nilotica and Kotpal (1971) on Penaeus lamarrez. The morphology and histology of the gut give ample evidence of the structural modifications of the gut to suit its food and mode of feeding. Paratya is seen to feed continuously for long intervals. The presence of a distensible oesophagus permits these quantities of food to be swallowed at a time. The oesophageal folds prevent regurgitation of food. Similar observations have been recorded in Panulirus (George et al., 1955), Metapenaeus bennettae (Dall, 1967) and Callzanassa (Powell, 1974). The stomach of crustaceans has been described as the gastric mill by various authors, for instance Parker (1876), Huxley (1880), Patwardhan (1935), Reddy (1935) , George et al. (1955) and Schaefer (1970). Review of the literature reveals in every case the stomach is differentiated into two regions, the anterior masticatory portion — the gastric mill — and a posterior pyloric portion — the filter apparatus. The size, shape and the structure of the gastric mill is variable in different groups. A strong gastric armature as identified by the presence of large calcified ossicles is mainly dependent on the size of the animal and the hard nature of the diet of shelled molluscs and arthropods. Though the word gastric mill is used by all the workers for its masticatory function the details given are highly variable. Paratya australiensis is a.small freshwater prawn feeding primarily on aquatic plants, algae, diatoms and insects. In association with the soft nature of the diet, and well-developed mandibles, the gastric mill in Paratya does not show any calcified ossicles as described by Huxley (1880), Patwardhan (1935) and Reddy (1935). The filter apparatus in the pyloric proventriculus is mainly used for sieving finer particles of food before they enter the midgut for digestion. The details of the filter apparatus show a strong resemblance to those observed in Cartdina laevis (Pillai, 1960), Corophium volutator and Caprella linearis (Aggarwal, 1963, 1964), Metapenaeus bennettae (Dall, 1967) and Callianassa (Powell, 1974). The pyloric brushes described by Powell were, however, absent in Paratya. The pyloric sheath in the mesenteron is a simple structure in Paratya and is functionally comparable to the terminal lappets of Penaeus setzferous (Young, 1959) and pyloric fingers of Callzanassa (Powell, 1974). Microscopic studies of the gut wall of Paratya show an inner layer of circular muscle fibres and an outer layer of longitudinal muscle fibres which are contrary to the observations of George et al. (1955) in Panulirus, and Vonk (1960) in Astacus. Pillai (1960) and Dall (1969) , however, observed the arrangement of muscle fibres in the gut wall of Carzdina laevis and Metapenaeus bennettae to be similar to that of Paratya. The digestive glands open into the mesenteron, which is the site for digestion and absorption of food. Intestinal caeca showing identical histological details to that of mesenteron can be inferred to perform similar functions. The present investigations are in agreement with the findings of George et al. (1955), Aggarwal (1963, 1964) Dall (1967) and Powell (1974). Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 216 AUSTRALIAN FRESHWATER PRAWN ACKNOWLEDGEMENTS I thank Professor D. T. Anderson, School of Biological Sciences, University of Sydney, Sydney, for his help in the preparation of this manuscript. My sincere thanks are due to Dr G. C. B. Poore, Senior Research Officer, Marine Studies Group, Victoria, for his helpful criticism of the manuscript. References AGGARWAL, V. P., 1963. — Functional morphology of the stomach of Corophium volutator. J. Linn. Soc. (Zool.), 45, No. 303: 47-52. ——, 1964. — Functional morphology of the stomach of Caprella linearis (L.). Current Science, No. 20: 618-619. CaLmaNn, W. T., 1909. — Crustacea. In Lankester’s Treatzse on Zoology, Vol. 7. London: Adam and Charles Black. Dati, W., 1967. — The functional anatomy of the digestive tract of a shrimp Metapenaeus bennettae Racek and Dall. Aust. J. Zool., 15: 699-714. Fryer, G., 1960. — The feeding mechanism of some atyid prawns of the genus Caridina. Trans. R. Soc. Edinb., 64: 217-244. GrorGE, C. J., RUBEN, N. and Mutu, R. T., 1955. — The digestive system of Panulcrus polyphagus (Herbst) . J. Anim. Morph. and Phys., 2: 14-27. Hux.ey, T. H., 1880. — The crayfish, an introduction to the study of Zoology. London: Kegan Paul, Trench, Trubner and Co., Ltd. KoTpaL, R. L., 1971. — Arthropoda. Meerut: Rastogi and Company Educational Publishers. Parker, T. J., 1876. — On the stomach of freshwater crayfish. J. Anat. Physzol., 11: 54-60. PATWARDHAN, S. S., 1934. — On the structure of the gastric mill in decapoda. Proc. Indzan Acad. Scz. B, 1: 359-375. ——, 1935. — The structure of the gastric mill in reptantons Macrura — Caridea. Proc. Indian Acad. Sct. B1: 414-422. PILLAI, S. R., 1960. — Studies on the shrimp Carzdzna laevis (Hellee). J. Mar. biol. Ass. India. 1 (1) : 57- 74, PoweELL, R. R., 1974. — The functional morphology of the foreguts of the thalassinid crustaceans, Callianassa californiensts and Upogebia pugettensts. Univ. California Publ. Zoology, 102: 1-47. Reppy, A. R., 1935. — The structure, mechanism and development of the gastric armature in stomatopoda with a discussion as to its evolution in Decapoda. Proc. Indian Acad. Sci., B1, 10: 650-675. SCHAEFER, N., 1970. — The functional morphology of the foregut of three species of decapod crustacea Cyclograpsus punctatus Milne Edwards, Diogenes brevirostris Stimpson and Upogebia africana Ortman. Zool. Africana, 5 (2) : 309-326. Vonk, H. J., 1960. — Digestion and Metabolism [in] The Physzology of Crustacea, Vol. I. New York and London: Academic Press. WiiiiaMs, W. D., 1977. — Some aspects of the ecology of Paratya australiensts (Crustacea, Decapoda: Atyidae). Aust. J. Mar. Freshwater Res., 28:403-415. YoncE, C. M., 1924. — Studies on the comparative physiology of digestion. II. Mechanism of feeding, digestion and assimilation in Nephros norvegicus. J. Exp. Biol., 1:343-389. Younc, J. M., 1959. — Morphology of the white shrimp, Penaeus setzferous (Linnaeus 1758). Fresh. Bull. U.S. Fish and Wildlife Service 59 (145) : 1-168. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 Three New Species of Amblysezus Berlese (Acarina: Phytoseiidae) from Australia E. SCHICHA ScHICHA, E., Three new species of Amblysezus Berlese (Acarina: Phytoseiidae) from Australia. Proc. Linn. Soc. N.S.W. 103 (4), (1978) 1979: 217-226. Adults and immature stages of Amblysezus lazlae n. sp. from papaw, adults of Amblyseius neolentiginosus n. sp. from Pinus radiata, and adults of Amblysezus neovictorzensis n. sp. from Indian coral tree (Erythrina sp.), all from coastal New South Wales, are described and illustrated. E. Schicha, Biological and Chemical Research Institute, N.S.W. Department of Agriculture, Rydalmere, Australia 2116; manuscript received 26 June 1978, accepted 20 September 1978. INTRODUCTION The three new species described here were collected in late summer 1978 during collecting trips along the coast of N.S.W. They are suspected of being predators of phytophagous mites. If not indicated otherwise, three specimens were measured of each of the develop- mental stages described, values being the range in micrometres (um). BCRI is the abbreviation for Biological and Chemical Research Institute, Rydalmere. Genus AMBLYSEIUS Berlese Amblysezus Berlese, 1914: 143. Type-species by original designation: Zercon obtusus Koch, 1839. Amblysezus lailae, n.sp. (Figs 1-22) Diagnosis. — In the adult stage, A. lazlae is similar to A. pafurtenses van der Merwe (1968) . However, in females of lazlae setae D4, M2, L3, L8 and L9 are slightly longer and setae D1 and L4 are considerably longer than those in pafurzenszs. In lazlae the macrosetae on genu III and tibia III are slightly shorter and on genu I, II and IV, tibia IV and basitarsus IV they are considerably shorter. The movable digit of the chelicerae in /azlae has three teeth, while that in pafurzenszs has only one. In lazlae the peritremes reach to the bases of D1 whereas in pafurzenszs they only reach to the bases of LI. Types. — NEW SOUTH WALES: on leaves of papaw, Alstonville, 18.iv.1978, M. Elshafie. Holotype 2 (A.laz.1) in BCRI; 4 99 (A.laz.2-5) and six dd (A.laz.6-7) paratypes; allin BCRI. Female (Figs 1-7) Dorsum. — Dorsal shield 343-372 long, 179-199 wide at L4, smooth, with 17 pairs of setae, six dorsal, two median, four prolateral, five postlateral: D1 30-32 long, D2 and D3 6-8, D4 8-9, D5 9-10, D6 7-8, M1 6-8, M2 9, L1 45-48, L2 8-10, L3 8-9, L4 63-71, L5 8-10, L6 11-12, L7 10-14, L8 7-10, L9 57-59. L9 slightly serrated, all other setae smooth. All setae shorter than distances between their bases and bases of setae following next in series, except for L1 which is longer than interspace L1/L2. Five pairs of large pores and two pairs of small pores as figured. $1 16-17 and S2 9-11 long. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 218 THREE NEW SPECIES OF AMBLYSEIUS BERLESE Figs 1-9. Amblysezus lailae, n.sp., female: 1, dorsum. 2, sternal shield. 3, ventrianal shield. 4, chelicera. 5, spermatheca. 6, leg III. 7, leg IV; male: 8, ventrianal shield. 9, spermatodactyl. The dimensions of these and other characters illustrated in Figs 1-38 are given in the text. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 E. SCHICHA 219 Peritremes extending forward to bases of D1 (Fig. 1). Venter. — Sternal shield 68-73 long, 84 wide, with three pairs of setae, two pairs of pores and lobate posteriorly. Fourth pair of setae on metasternal shields (Fig. 2). Vase-shaped ventrianal shield 116-121 long, 73-79 wide, with three pairs of preanal setae of equal length and one pair of preanal pores 34-37 apart (Fig. 3). Chelicera. — Fixed digit 33-37 long, with nine teeth plus pilus dentilis. Movable digit 37-39 long, with three backward pointing teeth (Fig. 4). Spermatheca. — Tube-like cervix 33-38 long, atrium occupying whole width where cervix fuses with macroduct, the latter very short (Fig. 5). Legs. — Seven macrosetae: spiniform on genu I 36 long, genu II 36-40, genu III 43- 44, tibia III 36-37 (Fig. 6) ; tapering apically and ending with a little knob on genu IV 73-77, tibia IV 61-62, basitarsus IV 106-111 (Fig. 7). Male (Figs 8-9) (two specimens measured) Dorsum. — Dorsal shield 280-290 long, 128-145 wide at L4, smooth, with chaetotaxy resembling that of female: D1 22-25 long, D2 to D4 4-5, D5 6-7, D6 5-6, M1 6, M2 8, L1] 35-36, L2 6, L3 8-9, L4 45-49, L5 to L7 8-10, L8 6-7, L9 43-46. S1 16 and S27, on interscutal membrane. Venter. — Imbricated ventrianal shield 114-117 long, 156-158 wide, with three pairs of preanal setae of equal length, the posterior two pairs on a transverse line, and a pair of pores 22-25 apart (Fig. 8). Spermatodactyl. — Shaft narrow; small foot broad with blunt toe and protruding heela(Fig=79))- Legs. — Seven macrosetae similar to those of female: on genu I 28, genu II 27, genu III 31, tibia III 27, genu IV 52-53, tibia IV 41-43, basitarsus IV 71-85. Deutonymph (Figs 10-13) Female (two specimens measured) Dorsum. — Smooth dorsal shield 296-319 long, 142-145 wide at L4, with 17 pairs of setae, six dorsal, two median, four prolateral, five postlateral: D1 28-30 long, D2 to D6 5-8, M1 6-7, M2 9, L1 43-44, L2 8, L3 8-10, L4 55-58, L5 11-12, L6 12-13, L7 12- 14, L8 6-10, L9 46-48. L9 slightly serrated, all other setae smooth. L1 as long as inter- space L1/L2; all other setae shorter than distances between their bases and bases of setae following next in series. Five pairs of large pores and three pairs of small pores as figured. S1 14-16 and S2 25 on interscutal membrane. Peritremes extending to L2 (Fig. 10). Venter. — Two pairs of preanal setae, two pairs of lateroventral setae, two pairs of posteroventral setae and one pair of caudal setae 32-33 long. Preanal pores 26-30 apart (Fig. 11). Chelicera. — Fixed digit 25-29 long, with 7-8 teeth and pilus dentilis. Movable digit 28-29 long with three backward pointing teeth (Fig.12). Legs. — Seven macrosetae similar to those of female (holotype): on genu I 35-36, genu II 36-37, genu III 48-49, tibia III 39-40, genu IV 67-71, tibia IV 51-61, basitarsus IV 94-95 (Fig.13). Protonymph (Figs 14-17) (one specimen measured) Dorsum. — Smooth dorsal surface 232 long, 122 wide at L4, bearing two shields. Anterior shield with nine pairs of setae, four dorsal, one median, four lateral; posterior shield with five pairs of setae, one dorsal, one median, three lateral; between Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 220 THREE NEW SPECIES OF AMBLYSEIUS BERLESE Figs 14-17. Amblysezus lailae, n.sp., protonymph: 14, dorsum. 15, venter. 16, chelicera. 17, leg IV. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 E. SCHICHA 221 the two shields three pairs of setae, one dorsal, two lateral: D1 20 long, D2-D6 3-7, M1 4,M29, L1 28, L2 and L3 7, L4 43, L5 to L8 5-9, L9 36. L9 slightly serrated, all other setae smooth. LI] as long as interspace L1/L2; all other setae shorter than distances between their bases and bases of setae following next in series. Peritremes 34 long (Fig. 14). Venter. — Two pairs of preanal setae, one pair of lateroventral setae, one pair of caudal setae, all of near equal length (Fig. 15). Chelicera. — Both digits 38 long. Fixed digit with six teeth and pilus dentilis; movable digit with three backward pointing teeth (Fig. 16). Legs. — Seven macrosetae similar to those of female (holotype) : on genu I 28, genu II 30, genu III 36, genu IV 65, tibia IV 62, basitarsus IV 78 (Fig. 17). Larva (Figs 18-22) (one specimen measured) Dorsum. — Smooth dorsal shield 218 long, 116 wide at L4. Ten pairs of smooth setae, four dorsal, one median, four prolateral, one caudolateral: D1 24 long, D2 and D3 3, DAViS el and 2713/8) 447 19 17 All setae shorter than distances between their bases and bases of setae following next in series. One pair of large pores between L4 and L9 and one pair of small pores near LO (Fig. 18). Venter. — Two pairs of preanal setae, posterior pair three times longer than anterior. Two pairs of lateroventral setae short. Preanal pores 19 apart (Fig. 19). Chelecera. — Both digits 21 long. Fixed digit with three teeth; movable digit with one tooth (Fig. 20). Legs. — Four macrosetae: on genu I 26, genu II 38, genu and tibia III 50 (Figs 21- i) 18 Figs 18-22. Amblysezus lailae, n.sp., larva: 18, dorsum. 19, venter. 20, chelicera. 21, leg II. 22, leg III. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 222 THREE NEW SPECIES OF AMBLYSEIUS BERLESE Amblysetus neolentiginosus, n.sp. (Figs 23-30) Diagnosis. — In the adult stage, A. neolentiginosus is similar to A. lentiginosus Denmark and Schicha (1975). However, in neolentiginosus setae D1 are shorter and setae M2, L1, L4, S1, S2 and all three macrosetae on leg IV are considerably shorter than those in lentzgznosus. In addition, in neolentiginosus setae L4 are only approx. half as long as those in lentzgznosus. In neolentigznosus the fixed digit of the chelicerae has 10 teeth whereas that in lentzgznosus has only seven teeth. Neolentiginosus has a well developed atrium between cervix and macroduct of the spermatheca, while lentegznosus has no distinct atrium. Types. — NEW SOUTH WALES: on Pinus radzata tips of branches, Narara, 2.ii1.1978, E. Schicha. Holotype Q@ (A.neol. 1); two QQ (A.neol. 2-3), one dD (A.neol. 4) paratypes; allin BCRI. Female (Figs 23-28) Dorsum. — Smooth dorsal shield 360-365 long, 177-183 wide at L4, with 17 pairs of setae, six dorsal, two median, four prolateral, five postlateral: D1 22-24 long, D2 to D6 4-8, M1 4-5, M2 77-78, L1 35-36, L2 and L3 7-10, L4 44-50, L5 to L8 4-7, L9 113- 115. M2 and L49 slightly serrated, all other setae smooth. L1 as long as interspace L1/L2, all other setae shorter than distances between their bases and bases of setae following next in series. Five pairs of large pores as figured. S1 and S2 4 long, on interscutal membrane. Peritremes extending forward beyond bases of D1 (Fig. 23). Venter. — Sternal shield 60-62 long, 75-80 wide, with three pairs of setae and two pairs of pores as figured. Fourth pair of setae on metasternal shields (Fig. 24) . Smooth pentagonal ventrianal shield 114-120 long, 94-100 wide, with three pairs of short preanal setae and a pair of preanal pores 17-19 apart (Fig. 25). Chelicera. — Both digits 33 long. Fixed digit with ten teeth plus pilus dentilis, movable digit with three teeth (Fig. 26). Spermatheca. — Tube-like cervix 16-18 long, atrium occupying whole width where cervix fuses with macroduct (Fig. 27). Legs. — Six macrosetae: on genu I 22-24, genu II 28-31, genu III 36-39, genu IV 53- 71, tibia IV 43-48, basitarsus IV 57-62 (Fig. 28). Male (Figs 29-30) (one specimen measured) Dorsum. — Dorsal shield 272 long, 156 wide at L4, with chaetotaxy resembling that of female, but all setae relatively shorter: D1 20 long, D2 to.D6 4-5, L1 31, L2 and L3 6, L4 37, L5 to L8 4-5, L9 90. S1 and S2 4 on interscutal membrane. Venter. Slightly creased ventrianal shield 114 long, 156 wide, with three short preanal setae, four pairs of small pores and a pair of large preanal pores 17-19 apart (Fig. 29). Spermatodactyl. — Shaft 17 long, foot ending with a knob (Fig. 30). Legs. — Six macrosetae similar to those of female: on genuI 19 long, genu II 21, genu III 23, genu IV 50, tibia IV 32, basitarsus IV 48. Amblysetus neovictoriensis, n.sp. (Figs 31-38) Diagnosis. — In the adult stage, A. neovictoriensis is similar to A. victoriensis (Womersley) (Womersley, 1954; see also Schicha, 1977). However, in neovictoriensis the dorsal shield is narrower and shorter, setae D1 are shorter, and setae L2 and L3 are approx. half as long as those in victoriensis. In neovictortensis setae L4 are longer Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 E. SCHICHA 223 Figs 23-30. Amblyseius neolentiginosus, n.sp., female: 23, dorsum. 24, sternal shield. 25, ventrianal shield. 26, chelicera. 27, spermatheca. 28, leg IV; male: 29, ventrianal shield. 30, spermatodactyl. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 224 THREE NEW SPECIES OF AMBLYSEIUS BERLESE Figs 31-38. Amblyseius neovictoriensis, n.sp., female: 31, dorsum. 32, sternal shield. 33, ventrianal shield. 34, chelicera. 35, spermatheca. 36, leg IV; male: 37, ventrianal shield. 38, spermatodactyl. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 E. SCHICHA 225 and setae, L6, L7 and L8 are approx. twice as long as those in wctorzenszs. The fixed digit of the chelicerae in neovectorzensis has 4 teeth, while that of vectorzenszs has 7 teeth. The two species differ in addition in the shape of their macrosetae of legs III and IV, their spermathecae and their spermatodactyli. Types. — NEW SOUTH WALES: on Indian coral tree (Erythrina sp.) , Coffs Harbour, 15.11.1978, E. Schicha. Holotype 9 (A.neo.1) in BCRI; two 29 (A.neo.2-3) and one dS (A.neo.4) paratypes all in BCRI. Female (Figs 31-36) Dorsum. — Dorsal shield 285-319 long, 182-186 wide at L4, reticulated antero- laterally, with 17 pairs of setae, six dorsal, two median, four prolateral, five post- lateral: D1 28-31 long, D2 to D6 6-8, M1 6-7, M2 9-11, L1 35-39, L2 33-34, L3 36-41, L4 43-46, L5 9-12, L6 17-21, L7 23-28, L8 23-31, L9 59-60. L9 slightly serrated, all other setae smooth. L1 longer than interspace L1/L2; L3 as long as interspace L3/L4. All other setae shorter than distances between their bases and bases of setae following next in series. Five pairs of pores as figured. S$] and S2 9-14, on interscutal membrane. Peritremes extending forward to base of L2 (Fig. 31). Venter. — Sternal shield 58-62 long, 71-76 wide, with three pairs of setae and two pairs of pores as figured. Fourth pair of setae on metasternal shields (Fig. 32). Vase- shaped ventrianal shield 89-94 long, 75-78 wide, with three pairs of long preanal setae and one pair of preanal pores 27-30 apart. Membrane surrounding ventrianal shield with three pairs of setae and four pairs of small shields. Caudolateral setae 32-37 long. Primary metapodal shield 11-14 long; secondary metapodal shield missing (Fig. 33). Chelicera. — Both digits 24-26 long. Fixed digit with four teeth plus pilus dentilis. Movable digit with one backward pointing tooth (Fig. 34). Spermatheca. — Sack-like cervix 11-13 long, atrium occupying whole width where cervix fuses with macroduct (Fig. 35). Legs. — Six macrosetae: spiniform on genu II 26-28 long, genu III 31-34, tibia III 28- 30, genu IV 46-51, tibia IV 36-43, basitarsus IV 63-71 (Fig. 36). Male (Figs 37-38) (one specimen measured) | Dorsum. — Dorsal shield 236 long, 148 wide at L4, with chaetotaxy resembling that of female: D1 26 long, D2 to D6 4-8, M1 6, M2 10, L1 32, L2 30, L3 31, L4 40, L5 8, L6é 17, L719, L8 26, L9 46. S1 and S2 14, on interscutal membrane. Venter. — Imbricated ventrianal shield 94 long, 156 wide, with three pairs of preanal setae, arranged in an almost straight line, and a pair of preanal pores 22 apart (Fig. 37). Spermatodactyl. — Shaft including foot 25 long; shaft narrow, pronounced heel rounded, toe ending with a suctorial disc (Fig. 38). Legs. — Six macrosetae similar to those of female: on genu II 19 long, genu III 26, tibia III 25, genu IV 37, tibia IV 33, basitarsus IV 48. ACKNOWLEDGEMENTS The assistance with the illustration of specimens by Mr M. Elshafie and Mrs J. Humphreys is acknowledged. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 226 THREE NEW SPECIES OF AMBLYSEIUS BERLESE References BERLESE, A., 1914. — Acari nuovi. Redza, 10: 113-150. DENMARK, H. A., and ScuicHA, E., 1975. — A new species of Amblysezus Berlese (Acarina: Phytoseiidae) from apple in Australia. Proc. Linn. Soc. N.S.W., 99: 145-150. Kocu, C. L., 1839. — Deutschlands Crustaceen, Myriapoden, und Arachniden. Regensburg. Merwe, G. G. van der, 1968. — A taxonomic study of the family Phytoseiidae (Acari) in South Africa, with contributions to the biology of two species. Ent. Mem. Dept. Agr. Techn. Serv. Sth Afr., 18: 1- 198. ScHICHA, E., 1977. — Amblysecus victoriensts (Womersley) and A. ovalis (Evans) compared with a new congener from Australia (Acari: Phytoseiidae) . J. Aust. ent. Soc., 16: 123-132. WomersLey, H., 1954. — Species of the subfamily Phytoseiinae (Acarina: Laelaptidae) from Australia. Aust. J. Zool., 2: 169-191. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 Biota of a Temperate Shallow Water Reef JEANETTE E. WATSON WATSON, J. E. Biota of a temperate shallow water reef. Proc. Linn. Soc. N.S.W. 103 (4), (1978) 1979: 227-235. This study describes the total flora and fauna of a small (1 m?) but representa- tive area of temperate water reef 23 m deep near Sydney, New South Wales. Excluding Foraminifera, the biota totalled 150 species comprising 29 algal, 120 invertebrate and 1 fish species. Hydroids are dominant in terms of species (13%) and sessile invertebrates comprise 51% of community biomass and 61% of species. Algae comprise 19% of species and 36% of community biomass. Populations of numerable animals are dominated by polychaetes (18%) and amphipods (17%). The sample yielded 8 new species, one new genus, extended the Australian provincial distribution of 17 species and provided 4 new records for Australia. Jeanette E. Watson, Honorary Assoczate, National Museum of Victoria, 285-321 Russell Street, Melbourne, Australia 3000; manuscript received 20 June 1978, revised manuscript accepted 18 October 1978. INTRODUCTION In recent years there have been a number of studies on the species composition, distribution and ecology of the soft bottom littoral and sublittoral estuarine com- munities of the south-eastern Australian coastline (King et al., 1971; Stephenson et al., 1974; Poore & Rainer, 1974; Hutchings & Recher, 1974). There are however, few published reports upon the epibiota of sublittoral reefs. These include several detailed studies of the ecology of benthic Algae in South Australian waters by Shepherd and Womersley (1970, 1971, 1976), and generalized accounts of the benthic ecology of Gulf St. Vincent and Investigator Strait in South Australia (Shepherd & Sprigg, 1976; Richardson & Watson, 1975a, 1975b), of Westernport Bay in Victoria (Smith et al., 1975), and of the central New South Wales coast near Sydney (Laxton, 1976, 1977; Jones, 1977. None of these studies has, however, been directed towards a complete analysis of the epibiota at any one sampling site. As part of a wider survey involving pollution studies at other localities on the central New South Wales coastline, the biota of a flat-topped sandstone reef in an unpolluted environment at 23 m depth was sampled at a site 500 m east of Jibbon Bombora in the ocean off Port Hacking (34° 01’ 20" S., 151° 14’ 20” E.) in September 1976. SAMPLING METHODS Following preliminary inspection, a site representative of the biota of the reef top was selected for sampling. A hoop of 0.33 m? was then thrown randomly on the site and the enclosed biota completely scraped into a fine mesh bag. This was repeated to give a total sampling area of 1 m’. The samples were later sorted under the microscope to species level and individual organisms counted. Colonial organisms were drained, blotted, and estimated as damp weight. This method of estimation of biomass was adopted since damp weighing does not destroy specimens which may later require detailed taxonomic examination. Only Foraminifera were not examined in detail. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 228 BIOTA OF A TEMPERATE SHALLOW WATER REEF RESULTS The list of species, together with numbers of individuals animals and the biomass of larger colonial organisms is given in the Appendix. Smaller colonial organisms are assigned a rank order of abundance from | (rare) ; 2 (common), to 3 (abundant). The biota of the total sampling area of 1 m* comprises 150 species, of which 29 (19%) are Algae belonging to 29 genera. One hundred and twenty species are inverte- brates, referable to at least 100 genera (not including sponges), and there is 1 species of fish. In terms of numbers of species, hydroids are the most abundant invertebrate group with 19 species (13% of total invertebrate species present) , followed by Bryozoa (15 species, 10%), polychaetes (15 species, 10%), amphipods (12 species, 8%), sponges (11 species, 7%), bivalves (10 species, 7%), and gastropods (8 species 5%). The collection contains 155 invididual non-colonial animals with the polychaetes con- tributing the greatest number (28 invididuals, 18%), followed by amphipods (27 individuals, 17%), gastropods (26 individuals, 17%), bivalves (25 individuals, 16%), and ophiuroids (17 individuals, 11%). The colonial polychaetes Phyllochaetopterus socialis, Serpula vermicularis and Filograna implexa, with relative abundances of (3), (2) and (1) respectively, would, if counted as individuals, considerably augment the polychaete population. Total biomass of Algae and animals amounts to some 700 g damp weight. In terms of biomass, the sponges, with 283 g (40% of biomass) are the most abundant group, followed by Algae with 255 g (36% of biomass), and the single alcyonarian colony, Capnella gaboensis, with 36 g (5% of biomass). The remaining biomass is made up of gastropods (3%), one species of ascidian, Polycarpa pedata (3%), bivalves and Bryozoa (each 2%) , and hydroids (1%). The more important groups of the biota are discussed below: Algae Of the 29 algal species recorded, 26 are red Algae and the remainder brown. All are small, low growing, turf-like species. Three are new records for New South Wales and include Antethamnion pinnafoltum Wollaston, previously unknown outside South Australia, Anotrichum crinitum (Kuetzing) Baldock, which has a recorded distribu- tion of southern Australia and New Zealand, and Schottera nicaeense (Duby) Guiry and Hollenberg, a Mediterranean species. Two as yet undescribed species, Propagula sp., and Medzothamnion sp., only recently discovered in Port Phillip Bay, Victoria, were also present in the sample. Several other species may also represent new taxa, but the material is insufficient for determination. Sponges This group comprises 11 species in 4 families. Only one, Callyspongia sp., a small grey dish-shaped sponge, could be identified to genus. The next most abundant sponge is an erect orange-coloured species belonging to the Raspailiidae. All are common sponges along this part of the New South Wales coastline. Hydroids The shallow water hydroid fauna of the New South Wales coast is known only from records of Bale (1884, 1888) and from the reports of the “Thetis” Expedition (Ritchie, 1911) and occasional incidental references in the literature. The 19 species of hydroids in the collection include 2 undescribed species, one belonging to the genus Hebella, and the other to Antennella. There are also 6 new records for the central east coast of Australia. Several of the newly recorded species are abundant in the sample and include the large plumose hydroid, Plumularia asymmetrica Bale, formerly recorded from the Great Australian Bight (Bale, 1915; Watson, 1973); Scoresbza daidala Watson, a small, obligatory epiphyte on the brown alga Zonarza crenata J. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 J. E. WATSON 229 Agardh, recorded once previously from the type locality in Gulf St. Vincent, South Australia, Monostaechus quadridens McCrady, formerly recorded from Queensland (Pennycuik, 1959), and Sertularia olsent Watson, a hydroid associated with sponges, recorded once, from the type locality in the Great Australian Bight (Watson, 1973). Only 2 of the hydroids, Tubularia australis Stechow, and Eudendrium sp., are athecate species. Of the 17 thecate species, 13 are small epiphytes growing among the algal turf, and the remainder, with the exception of the epilithic Plumularza asymmetrica, are epizooites, growing upon tubiculous polychaetes or ‘on other hydroids. Alcyonaria This group is represented by 3 species, including one large colony of Capnella gaboenszs Verseveldt. Although only recently described (Verseveldt, 1977), this is a common shallow water alcyonarian of New South Wales coastal reefs. Bryozoa The bryozoan fauna of the New South Wales coastline is not well known, and of the 12 species represented in the sample, 5 are new records for this part of the Australian coastline. There are 7 species of crustose (membraniporiform) Bryozoa, 4 erect jointed (cellariiform) species and 1 erect (vinculariiform) species. The cellariiform species, Crzsza acropora Busk and Emma triangula Hastings, are among the new locality records and are also the most abundant species in the sample. C. acropora is known from Bass Strait (Busk, 1875) and E. trzangula is known from this locality as well as from New Zealand (Hastings, 1939). Polychaetes Of the 15 species of polychaetes present, tubiculous forms are the dominant group in terms of numbers and include 5 species of serpulids. Phyllochaetopterus soczalis Claparéede is the most abundant polychaete in the sample (abundance (3) ), the clustered tubes providing substrate for a number of small sessile species, including hydroids and Bryozoa. The cosmopolitan Nematonerezs unicornis Grube is recorded for the first time in Australian waters. The nearest locality record for this species is the Indo-west Pacific. Approximately half of the polychaete species and the major part of the polychaete population are filter feeders, including all of the tubiculous species. The remainder may be either omnivores or predators but there is little information on their feeding habits. Mollusca The shelled molluscs are all small reef dwelling species well known from the literature. The sample comprises 26 gastropods belonging to 8 species and 25 individuals of 10 bivalve species. The predatory boring gastropod, Torvamurex denudatus (Perry) is the most abundant mollusc with 11 individuals in the sample. Since few bored bivalve shells were found, it seems likely that this species may be an unspecific predator or scavenger. The algal browsing seahare, A plysza sydneyensis Sowerby, is represented by one specimen. The type locality of this circumglobal species is Port Jackson. The remaining 2 species of opisthobranchs are undescribed species of the genera Phzline and Trapanza, the latter being a first record of this cosmopolitan genus for Australia. Amphipoda There are 2 undoubted, and possibly 5 additional undescribed species, reflecting the poor state of knowledge of the taxonomy of the group. Mallacoota subcarinata Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 230 BIOTA OF A TEMPERATE SHALLOW WATER REEF (Haswell), commonly occurring throughout south-eastern Australia, is the only species whose distribution is known. Half of the 11 amphipod species and the tanaid Paratanais cf. ¢gnotzs (Chilton) are tubiculous forms which are usually responsible for stabilization of substrate, accumulating silt and detritus in and around their tubes. Isopoda Of the 3 species of isopods recorded, only one, Cymodoce aspersa (Haswell) , whose type locality is Port Jackson, has been fully described. A single juvenile female is probably Czlicaeopsis granulata (Whitelegge) . Jibbon Head, adjacent to the sampling site, is one of the localities where this species was collected by Whitelegge (1902). The third species, represented in the sample by 2 adult males is an undescribed species which may also be worthy of separate generic status (W. Seed, pers. comm.). A similar, unnamed specimen was recorded from Masthead Island on the Great Barrier Reef by Baker (1926). Ophiurotdea The 4 species of ophiuroids include one new genus and one new record for the Aus- tralian east coast. The undescribed genus is an amphiurid related to the cosmopolitan shallow water species Amphipholis squamata (Delle Chiaje) (A. Baker, pers. comm.), and the new record is Ophiacantha alternata A. M. Clark, a small sub- littoral form previously known from Port Phillip Bay, Victoria. The 2 other species, Ophioactis restliens Lyman and Ophiothrix caespitosa Lyman are common com- ponents of the immediate sublittoral algal-sponge-bryozoan fauna of temperate Australia. Holothurotdea Of the 3 species identified, Pentacta australis Ludwig is a common form widely distributed along the Australian coastline. The other two species comprise Neocucumts ? wataset (Oshima) , known only from Japan, and an undescribed species of Thyone. Other Spectes The brachiopod Magellanza flavescens (Lamarck) represented by one small specimen, is a widely, but sparsely distributed species in shallow ocean waters of temperate Australia. Balanus trigonus (Darwin) is one of the commonest barnacles of the sublittoral of the New South Wales coastline, where it has been dredged from depths of up to 108 m (Pope, 1945). The ascidian Polycarpa pedata Herdman, is a very common solitary ascidian of the sublittoral of the south-eastern coast. It frequently grows in association with the stalked ascidian, Pyura spinzfera (Quoy & Gaimard) (J. E. W. unpub.). DISCUSSION The 150 species belonging to 104 genera include a new genus of ophiuroid and a probable new genus of isopod. Eight are probable new species (2 hydroids, 2 opisthobranchs, 2 amphipods, 1 isopod and 1 holothurian). The ranges of 4 algal species, 6 hydroids, 5 bryozoans and 1 ophiuroid are extended to the temperate east coast from the cool temperate southern region and the range of one isopod species is extended southwards from the Great Barrier Reef. There are 4 new records for Aus- tralia with the discovery of the cosmopolitan polychaete Nematonerevs unicornis, the holothurian Neocucumis ?watasez, previously recorded from Japan, the opistho- branch, Trapania sp., and the Mediterranean red alga, Schottera nicaeense. This number of new species and new records is all the more remarkable when the small size of the sampling area, and its situation in a region considered to be relatively well known scientifically, is taken into account. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 J. E. WATSON 231 With the exception of Callyspongza, the larger of which was 10 cm in diameter, and the ascidian Polycarpa pedata, the biota comprised small to very small organisms, most of which are dependent upon the algal turf for substrate, food and shelter. The algal community is dominated by small filamentous rhodophytic species; there are only 3 small species of brown Algae, and green Algae are absent. Sessile invertebrates comprise 51% of the biomass of the community and 61% of the species present. Algae account for 36% of the biomass and 19% of the species; algal cover was subjectively estimated at 50% of the sampling area. Analysis of community trophic structure shows that the greatest contribution to the animal community biomass (48%) is by the suspension feeding component, dominated by sponges. In terms of species, however, carnivores, mainly microcarni- vores, comprise the largest single feeding class with 42% of the total species present. Using photographic techniques in benthic studies off the New South Wales coastline, Laxton (1976, 1977) and Jones (1977), found that microcarnivores contributed the greatest percentage cover on most rocky substrates in areas where water turbidity is high. Pequengnat (1964), using species and population counts, found that suspension feeders were the most abundant components of a Californian reef-top habitat. Analysis of the present very limited sample supports Laxton’s findings; it is quite likely however, that habitat or geographical differences as well as sampling methods may lead to widely varying results. The new locality records from this sample provide further information upon the biogeographic affinities of south-eastern Australia with the Indo-Pacific and Japan. The sample also provides additional evidence for the close relationship of the Aus- tralian Peronian with the southern Flindersian Province west of Bass Strait. ACKNOWLEDGEMENTS The assistance of the following specialists who identified material and gave helpful advice and information is gratefully acknowledged: Dr. A. N. Baker, National Museum of New Zealand, Wellington, New Zealand (ophiuroids), Mr. R. Burn, Mr. D. A. Staples and Ms. S. Boyd, National Museum of Victoria, Melbourne (opisthobranchs, pycnogonids and bivalves respectively), Dr J. D. Kudenov and Dr G. C. B. Poore, Marine Studies Group, Ministry for Conservation, Melbourne (polychaetes and Crustacea), Messrs C. O’Brien and J. Lewis, Botany Department, University of Melbourne (Algae), Dr F. W. E. Rowe, Australian Museum, Sydney (holothurians) , Mr W. Seed, Department of Biology, Royal Melbourne Institute of Technology (isopods), Dr J. Verseveldt, Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands (Alcyonaria), and Dr A. Ayling and Mr R. Whitten, University of Auckland, New Zealand (sponges and Bryozoa). The author also wishes to thank Mr D. A. Staples and Mr H. Habgood for assistance with collection of material under difficult conditions. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 232 BIOTA OF A TEMPERATE SHALLOW WATER REEF References Baker, W. H., 1926. — Species of the isopod family Sphaeromidae from eastern, southern and western coasts of Australia. Trans. R. Soc. S. Aust., 50: 247-279. Bate, W. M., 1884. — Catalogue of Australian Hydrotd Zoophytes. Sydney: Australian Museum. ——, 1888. — On some new and rare hydroids in the Australian Museum collection. Proc. Linn. Soc. N.S.W., 3: 745-799. ——, 1915. — Report on the Hydroida collected in the Great Australian Bight and other localities. Part III. Bzol. Res. F.I.S. “Endeavour”, 3 (5) : 241-366. Busk, G., 1875. — Catalogue of the Cyclostomatous Polyzoa in the Collection of the British Museum. London: British Museum (Nat. Hist.). Ciark, A. M., 1966. — Port Phillip Survey 1957-1963. Echinodermata. Mem. Natn. Mus. Vict., 27: 289- 351. GruseE, A. E., 1840. — Actinien, Echinodermen und Wurmer des Adriatischen- und Mittelmeefes .. . KOnigsberg. Hastincs, A. B., 1939. — Notes on some cellularine Polyzoa (Bryozoa). Novitates Zoologicae, XLI: 321- 344. Haswe.u, W. A., 1881. — On some new Australian marine Isopoda. Proc. Linn. Soc. N.S.W., 5 (1): 470- 481. Hurcuincs, P. A., and Recuer, H. F., 1974. — The fauna of Careel Bay with some comments on the ecology of the mangrove and seagrass communities. Aust. Zool., 18 (2) : 99-128. Jones, A. R., (ed.), 1977. — An ecological survey of the nearshore waters east of Sydney, N.S.W. 1973-75. Sydney: Australian Museum. Kine, R. J., BLack, J. H., and Ducker, S. C., 1971. — Port Phillip Survey 2. Intertidal ecology of Port Phillip Bay with systematic list of plants and animals. Mem. Natn. Mus. Vict., 32: 93-128. LaxTon, J. H., 1976. — Ocean disposal of sewage effluent from coastal cities. A biologist’s viewpoint. Report to the Department of Public Works, New South Wales, Sydney. ——, 1977. — Boulder Bay Ocean Outfall. Report on Enwronmental Impact on marine communittes. Report for the Hunter District Water Board, Newcastle. Lyman, T., 1879. — Ophiuridae and Astrophytidae of the “Challenger” Expedition. II. Bull. Mus. Comp. Zool. Harvard, 6 (2) : 17-83. PENNYCUIK, P. R., 1959. — Faunistic records from Queensland. Part V. Marine and brackish water hydroids. Pap. Dep. Zool. Univ. Qld, 1 (6) : 141-210. PEQUEGNAT, W. E., 1964. — The epifauna of a Californian siltstone reef. Ecology, 45: 272-283. PooreE, G. C. B., and Ratner, S., 1974. — Distribution and abundance of soft bottom molluscs in Port Phillip Bay, Victoria, Australia. Aust. J. Mar. Freshwat. Res., 25: 371-411. Pope, ELIZABETH C., 1945. — A simplified key to the sessile barnacles found on the rocks, boats, wharf piles and other installations in Port Jackson and adjacent waters. Mem. Aust. Mus., 21 (6) : 351-372. RICHARDSON, JoYcE R., and WaTSON, JEANETTE E., 1975a. — Locomotory adaptations in a free-lying brachiopod. Science, 189: 381-382. —, 1975b. — Form and function in a recent free living brachiopod Magadina cumingz. Palaeobiology, 1: 379-387. RitcuiE, J., 1911. — Hydrozoa (Hydroid zoophytes and Stylasterina) of the “Thetis” Expedition. Mem. Aust. Mus., 4: 807-869. SHEPHERD, S. A., and Wome_rsLEy, H. S. B., 1970. — The sublittoral ecology of West Island, South Aus- tralia. 1. Environmental features and algal ecology. Trans. R. Soc. S. Aust., 94: 105-138. ——, 1971. — Pearson Island Expedition, 1969. 7. The subtidal ecology of benthic algae. Trans. R. Soc. S. Aust., 95 (3) ; 155-167. ——, 1976. — The subtidal algal and seagrass ecology of St. Francis Island, South Australia. Trans. R. Soc. S. Aust., 100 (4) : 177-191. SHEPHERD, S. A., and SpricG, R. C., 1976. — Substrate, sediments, and subtidal ecology of Gulf St. Vincent and Investigator Strait. Natural History of the Adelaide Region. Adelaide: Royal Society of South Australia Inc. Situ, B. J., CoLEMAN, N., and WaTSON, JEANETTE E., 1975. — The invertebrate fauna of Westernport Bay. Proc. R. Soc. Vict., 87 (1) : 149-156. Sowersy, G. B., 1869. — [Aplysza] Conchologia Iconica: XVII, Pl. 7, Fig. 31. London. STEPHENSON, W., WILLIAMS, W. T., and Cooke, S. D., 1974. — The benthic fauna of soft bottoms, southern Moreton Bay. Mem. Qld Mus., 17 (1) : 73-123. VERSEVELDT, J., 1977. — Australian Octocorallia (Coelenterata). Aust. J. Mar. Freshwat. Res., 28: 171- 240. WATSON, JEANETTE, E., 1973. — Pearson Island Expedition 1969. 9. Hydroids. Trans. R. Soc. S. Aust., 97 (3) : 153-200. WHITELEGGE, T., 1902. — Screntific results of the trawling expedition of H.M.C.S. “Thetis”. Crustacea Part III. Isopoda Part II. Sydney: Australian Museum. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 J. E. WATSON APPENDIX LIST OF SPECIES Numbers after the species name represent individual organisms. 233 Numbers in parentheses are an estimation of rank order of abundance of colonial organisms and algae, (1) =rare, (2) =common, (3) = abundant. New records are marked *. Probable new genera or new species are marked’. ALGAE Phaeophyta: Halopteris platycena Sauvageau (2) Dictyota alternifida J. Agardh (2) Zonaria sinclairz Hooker & Harvey (3) Rhodophyta: Delisea fimbriata (Lamouroux) Mont. (1) Cheilosporum sagittatum (Lamouroux) Aresch. (1) Jana sp. (1) Thamnoclonium sp. (1) Callophylilis sp. (1) *Schottera ncaeense (Duby) Guiry & Hollenberg (1) Glioderma sp. (1) Aglaothamnion sp. (1) *Anotrichium crinitum (Keutzing) Baldock (1) *Antithamnion pinnafolcum Wollaston (1) Balliella sp. (1) Callithamnion sp. (1) Ceramium sp. (3) *Mediothamnion sp. (3) Platythamnion sp. (3) *Propagula sp. (1) Pleonosporium sp. (1) Spongoclonium sp. (1) Acrosorium decumbens (J. Ag.) Kylin (1) Phycodris sp. (2) Dasya wilsonis J. Agardh (1) Heterostphonia australis (J. Ag.) De Toni (1) Aphanacladza sp. (1) Amplisiphona sp. (1) Dasyclonium incisum (J. Ag.) Kylin (1) Polystphonza sp. (1) INVERTEBRATA Porifera: Callyspongiidae Callyspongza sp. (2) Raspailiidae (3) Suberitidae (3) Spongiidae (2) Hydroida: Athecata: Tubularza australis Stechow (1) Eudendrium sp. (1) Thecata: *Clytza sp. (1) Obelza sp. (3) Hebella scandens (Bale) (1) *Hebella sp. (1) Hincksella cylindrica (Bale) (1) Halecium sessile Norman (1) *Halecium ? fragile Hodgson (1) Halecium delicatulum Coughtrey (1) *Scoresbia daidala Watson (2) Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 234 BIOTA OF A TEMPERATE SHALLOW WATER REEF Sertularella simplex Coughtrey *Amphisbetia olsent Watson Antennella secundaria Gmelin *Antennella sp. 2 *Monostaechus quadridens McGrady Plumularia spinulosa Bale Plumularia obliqua Saunders *Plumularia asymmetrica Bale Actinaria: Anemone sp. Alcyonaria: Capnella gaboensis Verseveldt Alcyonaria sp. 2 Alcyonaria sp. 3 Bryozoa: *Aetea anguina (Linnaeus) Chaperza cervicornis (Busk) Caberea helicina Hastings *Emma triangula (Hastings) *Celleporaria intermedia (MacGillivray) Celleporaria (Sinuporaria ?) hastigera Discopora vultur (Hincks) Smittotdea acaroensis (Busk) Celleporina sp. *Crisia acropora Busk Bicrista edwardsiana (d’Orbigny) *Pustulopora ? intricarta (Busk) “Retepora’ sp. Scuticella ventricosa (Busk) Amathia bisertata Krauss Polychaeta : Ophiodromus sp. Haplosyllis spongicola Grube Pionosyllis sp. Typosyllis sp. Exogone sp. Platynerets australis (Schmarda) *Nematonereis unicornis (Grube) Phyllochaetopterus socialis Claparéde Idanthyrsus pennatus (Peters) Branchiomma nigromaculata (Baird) Hydrordes ? brachycantha Rioja Pomatostegus polytrema (Philippi) Filograna implexa Berkley Serpula vermicularis Linnaeus Spirobranchus giganteus (Pallas) Sipunculida : Sipunculid sp. Nemertina: Nemertine sp. Brachiopoda: Magellania flavescens (Lamarck) Gastropoda: Torvamurex denudatus (Perry) Dentimitrella lincolnensis (Reeve) Cypraea sp. Notosinister maculosa (Hedley) Mitra carbonaria Swainson Austrosassia parkinsoniana (Perry) Vermicularza sp. Murexsul braziert (Angas) Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 (2) — — ss (1) (3) DS OWS ESS OS NES OS OS OS OS ESOS — DO DO DO KH 09 COR eS eS eS Oo Oe eee eee eee eee SS o——~S ee nwmoY Ope Horne —~ oa — ave — I yo) — —_ NmNMNMON fee J. E. WATSON 235 Ophisthobranchia: Aplysia sydneyensts Sowerby 1 *Philine sp. 1 *Trapania sp. 1 Bivalvia: Dosinia sp. Venerupts exotica Lamarck Mratella australis (Lamarck) Diplodonta globulosa A. Adams Marzkelllza solida (Angas) Cardita calyculata (Linnaeus) Atrina tasmanica (Tenison Woods) Lanistina impacta (Hermann) Lima nim bifer Iredale Trichomya hirsuta (Lamarck) Crustacea: Amphipoda: Photzs sp. Aora sp. Gammaropsis sp. Amphithoid sp. 1 Amphithoid sp. 2 Cerapus sp. *Phlianthid sp. *Liljeborgia sp. Mallacoota subcarinata (Haswell) Tethygenecza cf. elanora Barnard ? Corophiid sp. Amaryllis macrophthalmus (Haswell) Tanaidacea: Paratanais cf. ignotus (Chilton) Isopoda: Cymodoce aspersa (Haswell) *cf. Cymodoce sp. Cilicaeopsis granulata (Whitelegge) Brachyura: Plagusia chabrus (Linnaeus) Halicarcinus ovatus Stimpson Micippa tuberculosa (H. Milne Edwards) Leptomithrax sternocostulatus (H. Milne Edwards) Anomura: : Paguristes sulcatus Baker 1 Cirripedia: Balanus trigonus Darwin 4 Pycnogonida: Achela sp. 2 Nymphon molleria 1 Echinodermata: Ophiuroidea: Ophiothrix caespitosa Lyman 1 *Pholiostigma watsonae (Baker m.s.) *Ophiacantha alternata A. M. Clark Ophiactes restliens Lyman Holothuroidea : Pentacta doliolum (Pallas) 1 *Neocucumis ? watasez (Ohshima) 1 *Thyone sp. 1 Ascidiacea: Polycarpa pedata Herdman 1 NSN Phe Nw PH KH Oo — Phe DD — P — — —& PO OO — — — po Co — DO PISCES Gobiosocidae 1 Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 A New Species of Lemnadza (Crustacea: Conchostraca) from the Granite Belt in Southern Queensland and Northern New South Wales J. A. WEBB and G. D. BELL Communicated by A. RITCHIE Wess, J. A., & BELL, G. D. A new species of Lemnadza (Crustacea: Conchostraca) from the Granite Belt in southern Queensland and northern New South Wales. Proc. Linn. Soc. N.S.W. 103 (4), (1978) 1979: 237-245. Eulimnadia is synonymized with Lzmnadza, as the character separating the two, the presence or absence of a spine on the lower distal angle of the telson, is gradational and therefore unsatisfactory. Lamnadza urukhaz sp. nov. is distinguished from other species of the genus by the evenly curved dorsal margin and moderate size of its carapace (up to 6.7 mm by 4.3 mm), its few growth lines (maximum 10), small number of pairs of legs (15-16), two almost equal segments of the sixth endite in the first pair of claspers, and lack of a spine on the lower distal angle of the telson. On the basis of the variability noted in some morphological characters both by previous workers and in the present study, the Australian species Limnadza cygnorum and L. rivolenszs may be synonymous. J. A. Webb & G. D. Bell, Department of Geology and Mineralogy, University of Queensland, St Lucia, Australia 4067; manuscript rececved 19 June 1978, accepted 18 October 1978. INTRODUCTION The Queensland conchostracan fauna has been little studied ; of the 23 previously known Australian species, only 3 have been recorded from Queensland. These are Limnadia rivolensis (Brady 1886) and Caenestheria berney: (Gurney 1927) from the Longreach district (Gurney, 1927), and Cyclestheriza hislopi (Baird 1859) from Rock- hampton (Sars, 1887). In addition, in the Queensland Museum there are specimens from Stradbroke Island provisionally identified as Lynceus cf. tatez (Brady 1886), and from Cunnamulla labelled Cyzzcus cf. dictyon (Spencer and Hall 1896) . Thus it is of considerable interest that several populations of conchostracans collected from small rainwater pools in the Granite Belt of southern Queensland and northern New South Wales proved on close examination to be a new species of Limnadia. Genus Lzmnadza Brongniart Limnadia Brongniart 1820, p.84; Daday 1925, p.147; Ueno 1927, p.281; Brehm 1933, p.31; Straskraba 1965a, p.263. Eulemnadia Packard 1874, p.55; Sars 1895, p.14; Daday 1926, p.1; Barnard 1929, p.251; Mattox 1954, p.6. Paralimnadza Sars 1896, p.15; Daday 1925, p.146. Type Species. Limnadza lenticularis (Linnaeus 1761). Diagnosis. Umbo lacking, lines of growth restricted to marginal portion of shell (1.e. large larval valve), dorsal margin smooth; frontal organ on top of pyriform appendage; first antennae long, unsegmented (after Straskraba, 1965a). Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 238 A NEW SPECIES OF LIMNADIA Discussion. The differences between Lizmnadza and Eulimnadza have been treated differently by different authors. Packard (1874) erected Eulzmnadza for species that were distinguished from those in Limnadza by having a narrower shell with a straighter dorsal margin, fewer lines of growth, larger gills, shorter flabellae, and fewer pairs of legs. But, as Sars (1895) and Straskraba (1965a) pointed out, the size and shape of the shell and the number of growth lines can be very unreliable characters; they are closely related to environmental conditions and the age of the individual (Massal, 1954), and often cannot be used to differentiate between species. The validity of using the size of the gills has been questioned by Brehm (1933) and Daday (1925), who felt that this feature also could be strongly altered by external conditions. The flabellum length varies considerably between species, from shorter than to nearly twice as long as the leg to which it is attached. The number of pairs of legs shows a similar variability from 16 to 26. For these reasons Sars (1895) questioned the validity of Eulamnadza, although he considered that one feature could definitely be used to separate the 2 genera: no males were known for any species of Lzmnadza. Daday (1925) and Barnard (1929) thought this an insufficient criterion, as although only females have been recorded for several limnadiid species, in most cases this is probably due to the small number of specimens collected (L. lentzcularis is an exception). Daday instead proposed a single morpho- logical distinction: the presence (Eulémnadiza) or absence (Limnadiza) of a spine on the lower distal angle of the telson. Since there is a complete gradation between these two extremes (Fig. 1), this character is somewhat unsatisfactory, as Barnard (1929), Straskraba (1965a) and even Daday (1925) have noted. Daday (1925) felt that Eulimnadia and Limnadia should be subgenera but retained them as genera for “historical and practical reasons’. Ueno (1927) adopted the position that species with numerous growth lines and 12-13 segments in the flagellae of the second antennae should be Lzmnadza, whereas those with 4-6 growth lines and 9-10 antennae segments would be referred to Eulimnadia. However, Brehm (1933) found that the subdivisions in the second antennae were indistinct, difficult to count, and could vary from 8 to 12 in one individual. 7 d ax c Fig. 1. Telsons of (a) Limnadia lenticular’s (Linnaeus 1761), (b) L. révolens’s (Brady 1886), (c) L. victoriensts (Sayce 1903), (d) L. texana (Packard 1874), and (e) L. similis (Sars 1900), showing variation in shape of lower distal angle (indicated by arrows). From Daday (1925), Straskraba (1965a), Sayce (1903) , and Daday (1926). Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 J. A. WEBB ANDG. D. BELL 239 Mattox (1954) reaffirmed that the number of pairs of legs, the shape of the lower distal angle of the telson, and the number of segments in the second antennae could be used to separate Eulimnadza from Limnadia. Some species can be assigned to either genus, depending which of the above criteria are used. L. urukhaz sp. nov. would be referred to Lzmnadza on the basis of its telson shape, but the presence of males, and the small number of pairs of legs (15-16) and antennae segments (8-10) would place it in Eulzmnadza. Most authors have relied entirely on the outline of the lower distal angle of the telson. Although this appears to show little variability within species, the gradation illustrated by Fig. 1 indicates that it is not a satisfactory character for distinguishing genera. Therefore Eulzmnadza should be synonymized with Limnadza. Sars (1896) differentiated Paralimnadiza from Limnadiza on the almost straight dorsal margin in the male carapace of the former, its large number of growth lines (up to 30), and relatively small larval valve (as little as 25% of overall valve length). However, the dorsal margin and larval valve of the female shell are only slightly different from those of many other limnadiid species, some of which have more growth lines. Daday (1925) synonymized the 2 genera and this is followed here. Distribution. World-wide. Limnadia urukhaz, sp. nov. (Figs 2-25) Types. Queensland: Sow and Pigs, near Stanthorpe, 26.iii. 1978, G. Bell, 1 d (holotype, W7499) , 5 db, 92 (W7500) ; Mt. Norman, near Stanthorpe, 9.iv. 1978, J. Surridge, 10 d, 19 Q, 4 juveniles (W7502) ; Stanthorpe, Nov. 1929, H. Jarvis, 4d, 22 (W268) . New South Wales: Bald Rock, near Tenterfield, 26.111. 1978, A. Ewart, 7d, 12 (carapaces only) (W7501). All specimens housed in the Queensland Museum. Diagnosis. Bisexual; male and female carapaces similar, with evenly curved dorsal margins, large larval valves, and up to 10 growth lines, maximum size 6.7 mm by 4.3 mm; 15-16 pairs of legs; 2 segments in sixth endite on first pair of claspers almost equal in length; no spine on lower distal angle of telson. Male. Dorsal margin of carapace always smoothly curved (Figs 2-5); antero-dorsal and postero-dorsal corners usually obtuse and angular (Fig. 3), but occasionally rounded (Figs 2, 4); rarely slight concavity ventral to postero-dorsal angle (Fig. 5). Maximum carapace measurements 6.7 mm by 4.3 mm, with 10 growth lines present. Larval valve constitutes as little as 55% of overall carapace length. Elliptical shell gland (or muscle scar) always contained entirely within larval valve, and greatest dimension 20-35% of overall carapace length. Colour of carapace varies during ontogeny: juveniles with no growth lines trans- parent; adults dark brown on larval valve with yellowish growth segments, and clear spot close to ventral margin of larval valve (Fig. 4, dotted circle) , forming “window” through which claspers often visible. With increase in number of growth segments, clear spot increases in size and shifts ventrally (Fig. 5). Micro-ornament of shell very subdued punctate, although ridges between punctae sometimes reticulate, rarely showing indistinct radiating pattern. Basal stalk of pyriform appendage undifferentiated in immature individuals (Fig. 6) ; normally very short in adults (Figs 8, 9) , occasionally long (Fig. 7) . Eye lobe (prominence carrying paired eyes) close or immediately adjacent to frontal organ; angular projection on anterior side usually noticeable (Figs 6-8), occasionally very small and indistinct (Fig. 9b). Occipital notch commonly right-angled although obtuse in immature specimens (Fig. 6). Ocellus elongate triangular to tear-drop in shape, rarely sub-elliptical. Rostrum extends beyond ocellus for approximately length Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 A NEW SPECIES OF LIMNADIA S ~ (4), (1978) 1979 . Soc. N.S.W., 103 J. A. WEBB ANDG. D. BELL 241 of ocellus, with well-rounded termination, rarely pointed in immature individuals (Fig. 6). First antennae long, reaching second, third or fourth segment of second antennae, with 3 to 6 non-setose papillae on anterior side, Size and distinctness of papillae vary considerably, even within one individual (Figs 9a, b). Scape of second antenna extends slightly beyond rostrum; each flagella has 7 to 10 segments, often poorly differentiated towards tip and difficult to count. Flagellae normally almost equal in length, rarely one only half as long as the other (Fig. 9b). Fifteen to 16 pairs of legs, first 2 modified as claspers. Third endite of claspers varies in length, and projection below fourth endite (thumb) often indistinct (Fig. 10a). Sixth endite (subapical appendage) in first pair of claspers bipartite, apical segment slightly longer than basal (Figs 10a, 11). Tip carries short setae, but none at joint between segments. Apical segment of sixth endite in second pair of claspers much longer than basal (Fig. 10b) , otherwise the two pairs of claspers similar. Posterior 8 to 12 abdominal segments typically with 1 to 2 dorsal spines, but occasionally middle 5-8 segments each have up to 8 dorsal setae. Dorsal margin of telson armed with 10 to 15 spines; posterior spine longest and straight, curved, or protruding beyond posterior margin of telson (Figs 12-14). Other dorsal spines vary irregularly in size, even between two halves of telson of one individual. Forked filament between third and fifth spines from anterior end. Caudal claws as long as or slightly shorter than dorsal margin of telson, with few small setae and usually one small spine on dorsal surface. Lower distal angle of telson right-angled or slightly obtuse, and corner angular or slightly rounded. Female. Female carapace occasionally with more arched dorsal margin (Fig. 16) and smaller dimensions (maximum 6.2 mm by 4.1 mm) than male; otherwise very similar (Fig. 15), except lacks clear spot mentioned previously. Female head (Figs 17, 18) resembles that of male except occipital notch obtuse and rounded, becoming less obtuse in older specimens. Rostrum extends short distance past ocellus, termination rounded and right-angled. First antennae extend only to first or second segments of second antennae, , and have 2 to 3 poorly differentiated papillae (Fig. 18), or occasionally up to 5 well- marked papillae (Fig. 17). Second antennae, number of pairs of legs and dorsal modifications of abdominal segments same as for male. Ninth and tenth legs each possess long narrow flabellum (Fig. 19), up to 1.5 times as long as leg. Posterior spine on dorsal margin of telson generally protrudes beyond and inclined to posterior margin (Figs 20, 21) ; this is less common in males. Eggs 0.14 to 0.20 mm in diameter, covered with irregular or semi-regular pattern of ridges (Figs 25 a-c). Largest clutch carried by any female about 80. Juvenile. Shell of smallest individual collected transparent with no growth lines, measuring 2.3 mm by 1.3 mm; dorsal margin only slightly curved (Fig. 22). At least 12 pairs of legs differentiated (posterior legs difficult to count), 6 to 7 segments in flagella of second antennae, and 9 spines on dorsal margin of telson (Fig. 24). No Figs 2-14. 2. Carapace (3), W7502. 3. Carapace (3), W7499 (holotype). 4. Carapace (3), W7502, showing clear spot (dotted circle). 5. Carapace (6), W7501, showing clear spot. 6. Head (3), W7502. 7. Head (3), W7500. 8. Head (3), W7499 (holotype). 9a, b. Head (3) W7500, showing both first antennae. 10a, b. First and second claspers, respectively, W7502. 11. First clasper, W7499 (holotype). 12. Telson (dS), W268. 13. Telson (6), W7500. 14. Telson (6), W7499 (holotype). All scale lines represent 1 mm. For the sake of clarity, setae at joints between segments of second antennae have been omitted. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 242 A NEW SPECIES OF LIMNADIA 2S) Figs 15-25. 15. Carapace (Q), W7502. 16. Carapace (2), W7500. 17. Head (2), W7502. 18. Head (9), W7500. 19. Flabellum, W7502. 20. Telson (2), W7500. 21. Telson (9), W268. 22. Carapace (juvenile) , W7502. 23. Head (juvenile), W7502. 24. Telson (juvenile), W7502. 25a-c. Eggs, showing pattern of surface ridges. All scale lines represent 1 mm. For the sake of clarity, setae at joints between segments of second antennae have been omitted. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 J. A. WEBB AND G. D. BELL 243 flabellae, claspers, or eggs developed. Frontal organ well differentiated, although pyriform appendage lacks basal stalk (Fig. 23). On slightly larger specimen, still with no growth lines, 15 pairs of legs visible while other features the same. With one growth line, number of segments in second antennae 7 to 10. In specimens with 2 growth segments dorsal margin more curved, 12 dorsal spines on telson, pyriform appendage has basal stalk, and flabellae present in females. These specimens have adult complement of legs, second antennae segments, and telson spines. Individuals with 3 growth lines show claspers or eggs, indicating full sexual maturity. The previous descriptions of males and females were based on such specimens. Ontogenetic variation in some characters can still occur after the onset of sexual maturity, as indicated already for the occipital notch in both sexes and the rostrum in males. Discussion. Limnadia urukhai differs from species previously assigned to Eulimnadza in its lack of a spine on the lower distal angle of the telson. With regard to the other species of Limnadza, it is readily differentiated from L. stanleyana King 1855, since in the latter the male carapace has an almost straight dorsal margin, whereas that of the female is strongly curved; furthermore the larval valve of L. stanleyana constitutes as little as 20% of the overall shell length. L. grobbenz Daday 1925 and L. woltereckz Brehm 1933 are both very large, reaching 19 by 13 mm and 22.3 by 16.6 mm res- pectively, with up to 45 growth lines. The dorsal margin in both species displays a slight concavity absent in L. urukhaz. L. urukhai can be distinguished from the three Australian species L. cygnorum (Dakin 1914), L. revolensis, and L. badia (Wolf 1911) by its smaller number of legs (15-16 as against 18-20), and by the two almost equal segments of the sixth endite of its first clasper, where the other species have the apical segment twice as long as the basal or have 3 segments. L. lenticularis differs in that no males of this species have ever been described (despite extensive collections) , the female rostrum is very acute, and there are 22-24 abdominal segments. The work of several previous authors, notably Straskraba (1965a) , demonstrated that intraspecific variation in some characters is considerable, and the present study has enlarged on this. Putting this information together reveals that the following characters are not taxonomically useful for limnadiids: presence of angular projection on eye lobe, ocellus shape, number and distinctness of papillae on first antennae (particularly in females), relative lengths of flagella of second antennae, dorsal modifications of abdominal segments, presence of small spine halfway along caudal claw, and inclination of posterior spine on dorsal margin of telson to that margin. In addition, Brehm (1933) found that the number of segments in the flagella of the second antennae can range from 8 to 12 in one individual, and illustrations in Daday (1926) and Barnard (1929) show that setae at the joints of the sixth endite of the claspers may be absent or present. Figures in Sars (1895) reveal that in L. stanleyana the surface of eggs varies from ridged to subdued spinose, and Dakin (1914) found that the shell gland may or may not be confined to the larval valve. The following characters, although they show considerable variation, are thought to be useful if treated cautiously: prominence of projection below thumb of clasper, shape of rostrum and occipital notch, length of first antennae, number and relative lengths of joints of sixth endite of first clasper, number of pairs of legs, number of dorsal spines on telson, number of setae on caudal claw, shape of lower distal angle of telson, shape of dorsal margin and maximum dimensions and number of growth lines of carapace, and relative sizes of larval valve and shell gland. Ontogenetic variation in any of these must be taken into account. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 244 A NEW SPECIES OF LIMNADIA TABLE 1 Differences between Limnadza cygnorum and L. rivolensis L. cygnorum L. rivolensis frontal organ slightly elongated normal number of pairs of legs 18 20 spine halfway along caudal claw absent present projection below thumb of clasper prominent moderately well-marked number and relative lengths of 2, apical one twice as 3, subequal segments in sixth endite long as basal of first clasper surface of eggs subdued spinose spinose Of the other characters, the following may be important but too little is known of their variability: size and shape of frontal organ, length of flabellae, presence of setae on first antennae, and colour and micro-ornament of shell. Among the 24 Australian conchostracan species there are probably several synonymies, as suggested by Williams (1968) ; detailed studies of European and North American faunas have considerably reduced the number of species there (Straskraba, 1965 a, b, 1966; Sissom, 1968; Wiltshire, 1974). Applying the above knowledge of character variability to the Australian limnadiids reveals that L. cygnorum and L. rivolensts may be synonymous, as intimated by Glauert (1924). Table 1 lists the differences between them; the only significant one refers to the sixth endite of the first clasper. If the apical segment in L. cygnorum is in fact divided in half, a possibility admitted by Dakin (1914) in his original description, then the differences separating the species are within the limits of intraspecific variation and they are synonymous. A final decision must await re-examination of the original specimens, as the published descriptions omit certain features. Distribution. Probably throughout the Granite Belt of southern Qld., extending into at least the northern portion of the New England Tableland in N.S.W. Ecological notes. The specimens of L. urukhaz collected occurred in small (1 to 2m? area) shallow (4 to 5 cm deep) rainwater pools on bare granite outcrops. However, only the larger of these pools were occupied by conchostracans, in association with slugs and chironomiid insect larvae. The insect larvae appear to eat the soft parts of dead conchostracans, such that dead individuals left overnight had only their shells remaining in the morning. One population of L. urukhaz on Mt. Norman was bright green due to a heavy infestation of algae, particularly on the legs. This was identified as Characium spp. by Dr A. B. Cribb, Dept. Botany, University of Qld. In some pools there were large numbers of individuals, many vigorously copulating. L. urukhaz is an active species; it generally swims with the long dimension of the shell at about 45° to the horizontal and its head upwards, but can adopt any other position. At the surface it turns upside down and moves along venter upwards. When frightened or resting it buries itself sideways or venter upwards in the bottom sediment. Many specimens were observed feeding in the algae on the pool bottoms. Specimens at the Sow and Pigs were believed to be about 9 days old when collected, since the only rain for several months had fallen 10 days previously. They survived for 3 days after collection, when they had a maximum of 4 growth segments. Relative proportions of males and females varied considerably in different collec- tions, females making up 30% to 85% of the specimens. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 J. A. WEBB AND G. D. BELL 245 ACKNOWLEDGEMENTS Thanks are due to Tony Ewart, Jan Surridge and Greg Williamson for collecting specimens, and to Ron Monroe, Curator of Crustacea, Queensland Museum, for the loan of specimens.from that institution. References BARNARD, K. H., 1929. — Contributions to the crustacean fauna of South Africa. 10. A revision of the South African Branchiopoda (Phyllopoda). Ann. S. Afr. Mus., 29: 181-272. BREHM, V., 1933. — Phyllopoden, Mitteilungen von der Wallacea — Expedition Woltereck, 5. Zool. Anz., 104: 31-40. BRONGNIART, A., 1820. — Memoire sur le Lzmnadza, nouveau genre des Crustaces. Mem. Mus. natn. Hist. nat., Parts, 6: 83-93. Dapay de Dregs, E., 1925. — Monographie systematique des phyllopodes conchostraces. Troisiéme partie. Annls Sct. nat., ser. 10, 8: 143-184. ——, 1926. — Monographie systematique des phyllopodes conchostraces. Troisiéme partie, suite. Annis Sct. nat., ser. 10, 9: 1-81. Dakin, W. J., 1914. — Fauna of Western Australia. II. The Phyllopoda of Western Australia. Proc. zool. Soc. Lond., 1914: 293-305. GLAUERT, L., 1924. — Contributions to the fauna of Western Australia. 5. Crustacea. J. Proc. R. Soc. West. Aust., 10: 59-64. Gurney, R., 1927. — Some Australian freshwater Entomostraca reared from dried mud. Proc. zool. Soc. Lond., 1927: 59-79. MassAL, L., 1954. — Deuxiéme note sur le milieu et la croissance des Estheries. Bull. Soc. Sct. nat. Tunis, 7: 165-181. Matrox, N. T., 1954. — A new Eulimnadiza from the rice fields of Arkansas with a key to the American species of the genus. Tulane Stud. Zool., 2: 3-10. PACKARD, A. S., 1874. — Descriptions of new North American Phyllopoda. Ann. Rep. Peabody Acad. Scu., 6. (vide Sars, 1895). Sars, G. O., 1887. — On Cyclestheria hislopi (Baird), a new generic type of bivalve phyllopod raised from Australian mud (Queensland). Vidensk. Selsk. Forh. Christ., 1: 1-65. —— , 1895. — Descriptions of some Australian Phyllopoda. Arch. Math. Naturv., 17: 1-51. ——, 1896. — Description of two new Phyllopoda from North Australia. Arch. Math. Naturv., 18: 1-36. Sayce, O. A., 1903. — The Phyllopoda of Australia, including descriptions of some new genera and species. Proc. R. Soc. Vict., 15; 224-261. Sissom, S. L. A., 1968. — A taxonomic review of the North American species of Eulizmnadza (Conchostraca, Crustacea) (abstr.). Dzss. Abstr., 28B: 4357. STRASKRABA, M., 1965a. — Taxonomic studies on Czechoslovak Conchostraca. I. Family Limnadiidae. Crustaceana, 9: 263-273. —, 1965b. — Taxonomic studies on Czechoslovak Conchostraca. II. Families Lynceidae and Cyzicidae. Vest. csl. Spol. zool., 29: 205-214. ——, 1966. ~— Taxonomic studies on Czechoslovak Conchostraca III. Family Leptestheriidae. Hydro biologia, 27: 571-589. Ueno, M., 1927. — Freshwater Branchiopoda of Japan. 1. Mem. Coll. Sct. Kyoto Univ., B2: 259-311. Witiiams, W. D., 1968. — Australian freshwater life. The invertebrates of Australian inland waters. Melbourne: Sun Books. WILTSHIRE, C. T., 1974. — The developmental morphology of Cyzécus morsez (Packard) (Crustacea: Conchostraca) from hatching through adulthood with comments on taxonomy within the family Cyzicidae. Diss. Abstr., 35B: 1132. Wo Lr, E., 1911. — Phyllopoda. In Die Fauna Sudwest-Australiens, 3: 253-276. W. Michaelsen and R. Hartmeyer (eds) . Jena: G. Fischer. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 te ‘i i Records of Little- Known Sharks from Australian Waters A.J. BASS Communicated by J. R. PAXTON Bass, A. J. Records of little-known sharks from Australian waters. Proc. Linn. Soc. N.S.W. 103 (4), (1978) 1979: 247-254. Chlamydoselachus anguineus, Hemipristis elongatus, Cirrhigaleus barbifer, Centroscymnus crepidater and Centroscymnus owstonz are recorded from Australia for the first time, illustrated and briefly described. A further record of Hexanchus griseus is noted. A. J. Bass, National Museum of New Zealand, Private Bag, Wellington, New Zealand; manuscript received 22 March 1978, accepted in revised form 13 December 1978. INTRODUCTION The shark fauna of Australia is not well known and many species often go unrecognized due to a lack of readily available descriptions. This paper attempts a partial remedy by recording and describing several obscure but possibly quite abundant species. Terminology and methods are as described in Bass et al. (1976). All study material is in the collections of the Australian Museum, Sydney (AMS) and the Western Australian Museum, Perth (WAM). Family CHLAMYDOSELACHIDAE Chlamydoselachus anguineus Garman 1884 Chlamydoselachus anguineus Garman 1884: 47, figs (type locality: Japanese seas) . Study material: 1315 mm mature male (AMS I.19157-001) trawled from 512-585 m of water off Brush Island, New South Wales (35°35-29'S, 150°44-47E), F.R.V. “Kapala”, 9 June 1976. This specimen is illustrated (Fig. 1) and its proportional dimensions summarized (Table 1). A vertebral count was not attempted as the lack of calcification in Chlamydoselachus renders most of the centra invisible in radiographs. The teeth number 13-13/12-1-12. Distinguishing features include the blunt snout, elongate body, single dorsal fin, anal fin, distinctive tricuspid teeth which are similar in the upper and lower jaws, and six gill-slits of which the first pair are joined across the throat. Fig. 1. 1315 mm mature male Chlamydoselachus anguineus (AMS 1.19158-001). A — lateral view, B — tooth from near centre of lower jaws. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 248 LITTLE-KNOWN SHARKS Stead (1907) recorded Chlamydoselachus from New South Wales on the basis of a skull with part of a vertebral column measuring over three metres in length. I agree with Whitley (1940) that this identification is highly doubtful. First described from Japanese seas where it is relatively common, C. anguzneus has also been caught in the north-eastern Atlantic on several occasions and twice off California (Roedel and Ripley, 1950). The only definite records from the southern hemisphere are of two sharks trawled at different times off South West Africa (Smith, 1967; Bass et al., 1975b) . To these may now be added the record from New South Wales. Family HEX ANCHIDAE Hexanchus griseus (Bonnaterre 1788) Squalus griseus Bonnaterre 1788: 663 (type locality: Mediterranean) . Study material: Jaws of a 4250 mm, 348 kg female (AMS I.19110-001) caught on a bottom long-line in 420 m of water off Norah Head, New South Wales (33°15'S, 152°15 E), J. Dulhunty, 6 June 1976. No detailed measurements of this shark are available as only the jaws were preserved. The tooth count is 18-19/9-VI-1-VI-8, the Roman numerals referring to the large multicuspid teeth found in the lower jaws of all hexanchid sharks. This group is distinguished by the presence of six or seven gill-slits (all widely separated across the throat), an anal fin, a single dorsal fin, and markedly different teeth in the upper and lower jaws. Of the four currently accepted species, Notorynchus cepedzanus (Peron 1807) and Heptranchias perlo (Bonnaterre 1788) are relatively common and well known in Australian waters. Heptranchias dakini Whitley 1931 is a synonym of H. perlo (see Garrick and Paul, 197la). The third species, Hexanchus griseus (Bonnaterre 1788), was first noted in Australian waters by Stead (1963) and then described and figured by Lynch (1964) on the basis of a 219 cm male caught in Victoria. The present study material constitutes the second definite record of H. griseus from Australia, for some of Stead’s specimens may have belonged to the fourth species, Hexanchus witulus Springer and Waller 1969. A demersal shark of warm waters, H. vitulus has been recorded in the western Atlantic — Gulf of Mexico region, the south-western Indian Ocean and the Philippines (see Bass et al., 1975b). In case of doubt the following key should serve to identify all four hexanchid species. Key to species of the family HEXANCHIDAE 1 SUXNGTITESINES | ene ueeely we cee aceite re nice) eNO a WANN acacia rouse te 2 — Seven gill-slits ; Lower jaws with five rows of large teeth on each side; dorsal fin height about twice anal height; mouth width about 1144 X mouth length Bin PH cM res Ee ee Rao Et A Sea OA SHA TER ON yee es AN na a Hexanchus vitulus — Lower jaws with six rows of large teeth on each side; dorsal fin height about 1144 x anal height; mouth width more than twice mouth length Bet Ry Rea irr ere Onn ERROR CMU Gm treacle MAN AU AA MeN A Hexanchus griseus 3 Snout sharp, its length more than 144 X distance between nostrils; body plain, without any small dark or white spots RUE ec Eel me A SI a STAIR ty eM ll gS SN Ue ME OAC ah Heptranchias perlo — Snout blunt, its length much less than 14% X distance between nostrils; upper surface of body with numerous small dark or white spots SESE Ne a eihad Say ces eae octet as EE, ae ame e sa ge Notorynchus cepedianus bo Family CARCHARHINIDAE Hemipristzs elongatus (Klunzinger 1871) Dirrhizodon elongatus Klunzinger 1871: 665 (type locality: Red Sea). Proc. LINN. Soc..N.S.W., 103 (4), (1978) 1979 A. J. BASS 249 Study material: 656 mm immature male (WAM P.24547) from Exmouth Gulf, Western Australia (22°10'S, 114°20’E), D. Heald, 6 July 1973; head and one pelvic fin of 1250 mm mature male (AMS I.19438-005) taken in a gill-net in 1-6 m of water off Lizard Island, Queensland (14°40'S, 145°27'E), Australian Museum party, 1-12 November 1975. Fig. 2. 656 mm immature male Hemzprist7s elongatus (WAM P.24547). A — lateral view, B —ventral view, C — denticle from side below first dorsal fin, D — teeth from right side of upper jaws. Dashed line indicates position of symphyses ; numbers indicate position of tooth rows counting from centre of jaws. The smaller of these two specimens is illustrated (Fig. 2) and its proportional dimensions summarized (Table 1). Vertebrae number 51 monospondylous in 101 precaudal and a total of 187. The teeth number 14-14/17-17 (13-13/16-16 in the larger shark). Colour (in alcohol) is a pale greyish brown, lighter ventrally, with no conspicuous markings on the fins. The head of the mature male (also in alcohol) is dark grey dorsally, pale cream below. H. elongatus is usually described as having spiracles which is not the case with the present study specimens. An interdorsal ridge is variously described as present or not (see Bass et al., 1975a). A definite ridge is present on the anterior half of the interdorsal space of the smaller of the two Aus- tralian sharks. Identification is based on the following suite of characters: distinctive teeth (Fig. 2D); definite upper and lower lip grooves; internal nictitating lower eyelids; first dorsal origin approximately over inner pectoral corner ; second dorsal fin about half as high as first dorsal, its origin anterior to the anal fin origin; anal fin slightly shorter than second dorsal fin; caudal peduncle without lateral ridges; precaudal pits present (the lower pit may be faint) ; caudal fin with a distinct lower lobe and subterminal notch; and falcate pectoral and pelvic fins, especially the latter which have acutely pointed tips to the lateral lobes.. The Australian fauna includes a number of carcharhinid sharks, many of them poorly known and usually not identified correctly. The specimens described above constitute the first record of Hemzprist7s elongatus from Australia. Previous records range from southern Africa and Madagascar north to the Red Sea and east to Vietnam. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 250 Summary of proportional dimensions, shown as percentages of total length Snout to outer end nostrils inner end nostrils mouth eye spiracle Ist gill-slit pectoral origin 1st dorsal origin pelvic origin upper caudal origin Horizontal eye diameter Spiracle length Upper ends Ist to last gill-slits 1st to 2nd dorsal origins Pectoral to pelvic origins Pelvic to anal origins Pelvic to lower caudal origins Anal to lower caudal origins Last dorsal to upper caudal origins Nostril length Distance between inner ends nostrils Mouth width Mouth length Upper lip groove ~ Lower lip groove Ist gill-slit 3rd gill-slit Last gill-slit 1st dorsal height base posterior lobe 2nd dorsal height base posterior lobe Anal height base posterior lobe Pectoral base inner edge length Pelvic anterior edge origin to tip Caudal upper lobe lower lobe Subterminal notch to caudal tip LITTLE-KNOWN SHARKS TABLE l C.anguineus H.elongatus 1315mm male 656mm male 0.8 3.8 0.9 4.0 0.2 7.2 2.1 6.3 8.0 16.3 14.7 21.5 61.3 29.0 48.5 49.5 76.4 76.5 1.4 2.4 6.7 5.5 - 30.5 33.7 25.9 12.2 17.1 13.2 14.2 15.1 17.1 0.7 1.5 3.3 4.0 7.6 5.6 6.4 4.1 = 1.4 = 0.9 6.3 4.1 5.4 4.0 4.3 2.9 2.6 9.3 10.0 10.4 2.6 2.6 = 4.4 = 32 -= 2.3 42, 3.5 12.6 6.4 1.4 2.4 3.5 4.7 a2 4.6 8.2 15.9 6.8 9.5 12.8 8.2 23.8 29.4 8.3 11.6 4.4 7.0 Family SQUALIDAE C.barbifer 704mm female 2.7 Cirrhigaleus barbifer Tanaka 1912 Cirrhigaleus barbifer Tanaka 1912: 151, pl. 141, figs 156-162 (type locality: Tokyo market, Japan). C.crepidater 338mm male 3.6 5.0 14.5 10.7 18.3 24.3 27.8 34.6 58.3 77.5 5.9 2.1 C.owstoni 975mm female Study material: 704 mm female (AMS I.19154-001) trawled from 494 m of water off Brush Island, New South Wales (35°34'S, 150°45-46E), F.R.V. “Kapala”, 6 July 1976. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 A. J. BASS 251 This shark is illustrated (Fig. 3) with the proportional dimensions summarized (Table 1). Definite keels are present on the caudal peduncle while the caudal fin lacks any trace of a subterminal notch. No marked precaudal pits are present but a faint shallow notch at the upper caudal origin could be construed as an incipient (or vestigial) precaudal pit. Vertebrae number 50 monospondylous in 85 precaudal and a total of 114. The teeth number 13-14/12-12. Colour (when freshly dead) dark grey above, white below, the fins with white trailing edges accentuated by somewhat darker colouring adjacent to the white edges. C. barbifer, distinguished by extremely long nasal barbels and by a lack of precaudal pits, was known only from two Japanese specimens until Garrick and Paul (1971b) recorded three from New Zealand and noted the existence of at least four others from Japan. The example from off Brush Island represents the first record from Australia. Garrick and Paul (1971b) commented on the close similarities between Squalus and Cirrhzgaleus but regarded the latter as valid. Bass et al. (1976) pointed out that the recently described Squalus asper Merrett 1973 bridges the gap between the two genera and concluded that they should probably be merged. For the present, however, I retain the genus Czrrhzgaleus while emphasizing its close relationship with Squalus asper. Fig. 3. 704 mm female Czrrhigaleus barbifer (AMS 1.19154-001). A — lateral view, B — ventral view, C — denticle from side below first dorsal fin, D — teeth from right side of upper jaws. Dashed line indicates position of symphyses. Genus Centroscymnus Bocage and Capello 1864 Four genera of Squalinae have laterally grooved fin spines and unicuspid teeth that are dissimilar in the upper and lower jaws. They are best distinguished by the characters used in the following key (after Bass et al., 1976). Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 252 LITTLE-KNOWN SHARKS Key to some genera of the subfamily SQUALINAE 1 Distance from snout to mouth longer than that from mouth to pectoral OUI IN SiiRebd ics istt tues nenshly canta amma BAe) GRA Deaniza Jordan & Snyder 1902 — Distance from snout to mouth equal to or less than that from mouth to pectoral OPIQDM sce eta ee tates BEE CS ea Sate ans Lt oe Goa al taal ean Re SUT NN Seagc NN Len 2 2 Upper teeth set close together, at least some of the bases overlapping to form an Intenlockangdb and) chee wee ae eee Centrophorus Muller & Henle 1837 — Upper teeth set apart, the bases not overlapping to form an interlocking band ARES GET are COUR Oe ene EM, AUT ET CULL cr ean UNM ACA an NUE Uae MS te WS Tyla bagi) A) 3 3 Upper teeth midway along either side of upper jaws distinctly longer than those towards centre; denticles of adults with a long median ridge, without a con- Spicuous median cavity, ..2065).5 540.148 Scymnodon Bocage & Capello 1864 — Upper teeth midway along either side of upper jaws not distinctly longer than those towards centre; denticles with median ridge absent or restricted to posterior part, with a conspicuous median concavity Uri eennue serail sete irie Spanair bina Centroscymnus Bocage & Capello 1864 Genus Centroscymnus Bocage and Capello 1864 Centroscymnus, as defined above, was not known from the southern hemisphere until Garrick (1959a) recorded two species in New Zealand. Centroscymnus crepidater (Bocage and Capello 1864) has since been taken off Namibia (Pinchuk and Permitin, 1970) while C. owstoni Garman 1906 has been caught in the south- western Indian Ocean (Forster et al., 1970). The occurrence of these two sharks in Australian waters is therefore not unexpected. The species are distinguished according to the following key. Key to Australian species of the genus Centroscymnus 1 Preoral clefts almost meeting in the midline of the upper jaws ... C. crepidater — Preoral clefts short, the distance between their inner ends at least as great as the distance between the inner ends of the nostrils ................. C. owstont Centroscymnus crepidater (Bocage and Capello 1864) Centrophorus crepidater Bocage and Capello 1864: 260 (type locality: Portugal) . Study material: 305 mm immature male (AMS I.15987-013) trawled from 549 m of water 48 km north-east of Jervis Bay, New South Wales (34°40°-35°01'S, 151°10- 07’E), F.R.V. “Kapala”, 7 July 1971; 338 mm immature male (AMS I.17868-003) trawled from 777 m off Sydney, New South Wales (33°40-43’S, 151°56-59'E), F.R.V. “Kapala’, 6 December 1972. The larger of these two specimens is illustrated here (Fig. 4) with the propor- tional.dimensions summarized (Table 1). The colour (in alcohol) is a uniform dark brown. The denticles are typical of young Centroscymnus; larger sharks have denticles similar to those illustrated for C. owstonz (Fig. 5C, see also Garrick, 1959a, fig. 4). Both specimens had 75 precaudal vertebrae with 54 (338 mm male) and 55 (305 mm male) monospondylous vertebrae. The caudal vertebrae were insufficiently calcified to make clear images on radiographs. Tooth numbers were 36/16-1-16 (338 mm male) and 37/16-1-16 (305 mm male). Centroscymnus owstoni Garman 1906 Centroscymnus owstont Garman 1906: 207 (type locality: Japan). Study material: 440 mm female (AMS IB.5327) trawled from 823 m of water east of Eden, New South Wales (37°04'S, 149°55’E), J. Henry, 29 August 1961 (in poor Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 A.J. BASS 253 ive Fig. 4. 338 mm immature male Centroscymnus crepidater (AMS I.17868-003). A — lateral view, B — ventral view, C — denticle from side below first dorsal fin, D — teeth from right side of upper jaws. Dashed line indicates position of symphyses. Fig. 5. 975 mm mature female Centroscymnus owstont (AMS 1.16147-001). A — lateral view, B — ventral view, C — denticle from side below first dorsal fin, D — teeth from right side of upper jaws. Dashed line indicates position of symphyses. condition) ; 975 mm mature female (AMS 1.16147-001) trawled off Sydney, New South Wales (151°E, 33°S), F.R.V. “Kapala”, August 1971. The 975 mm female is illustrated (Fig. 5) with the proportional dimensions sum- marized (Table 1). Many of the differences in proportions between this mature shark and the immature C. crepzdater illustrated in Fig. 4 are due to growth changes rather than specific differences. Trends in the changes of proportional dimensions with growth in squaloid sharks are summarized by Garrick (1960). The colour (in alcohol) is a uniform dark brown. Vertebrae numbered 57 monospondylous in a precaudal total of 75 (caudal vertebrae not counted). The teeth numbered 37/17-1-17. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 254 LITTLE-KNOWN SHARKS The smaller of these two specimens of C. owstonz was identified as Scymnodon plunketi Waite 1910 by Whitley (as noted in the Australian Museum fish register) . S. plunketz had previously been recorded from south-eastern Australia by Cowper and Downie (1957) together with Centroscymnus waited (Thompson 1930). Garrick (1959b) has since shown that C. watte7 is the juvenile of S. plunket?. The specimens noted by Cowper and Downie included Centrophorus squamosus (Bonnaterre 1788), Centroscymnus owstoni and one shark which probably was S. plunket: (pers. comm., J. A. F. Garrick). S. plunketz could well appear in catches of deepwater sharks from Australian seas. Apart from the tooth and denticle characters noted in the generic key above, it is readily distinguished from C. crepidater and C. owstonz by a shorter snout tip (distance between inner ends nostrils about 14% X that from tip of snout to inner ends nostrils in S. plunketz, about equal in C. crepzdater and C. owstonz2) . ACKNOWLEDGEMENTS For specimens, help, advice and hospitality I would like to thank Dr John Paxton, Dr Doug Hoese, Dianne Blake and Helen Larson of the Australian Museum as well as Dr Gerry Allen, Barry Hutchins and Neil Sarti of the Western Australian Museum and Dr Jack Garrick of the Victoria University of Wellington. Many of the specimens were obtained through the good services of Terry Gorman, Ken Graham and the crew of the F.R.V. “Kapala” of the New South Wales State Fisheries. References Bass, A. J., D’AuBREy, D. D., and KistNasamy, N., 1975a. — Sharks of the east coast of southern Africa. III. The families Carcharhinidae (excluding Mustelus and Carcharhinus) and Sphyrnidae. Invest. Rep. oceanogr. Res. Inst., (38) : 1-100. ——, 1975b. — Sharks of the east coast of southern Africa. V. The families Hexanchidae, Chlamydo- selachidae, Heterodontidae, Pristiophoridae and Squatinidae. Invest. Rep. oceanogr. Res. Inst., (43) : 1-50. ——, 1976. — Sharks of the east coast of southern Africa. VI. The families Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. Invest. Rep. oceanogr. Res. Inst., (45) : 1-103. Cowper, T. R., and Downir, R. J., 1957. — A line-fishing survey of the fishes of south-eastern Australian continental slope. C.S.I.R.O. Aust. Div. Fish. oceanogr. Rep., (6) : 1-19. Forster, G. R., and collaborators, 1970. — Results of the Royal Society Indian Ucean Deep Slope Fishing Expedition, 1969. Proc. R. Soc. Lond., B. 175: 367-404. Garrick, J. A. F., 1959a. — Studies on New Zealand Elasmobranchii. Part VIII — Two northern hemi- sphere species of Centroscymnus in New Zealand waters. Trans. R. Soc. N.Z., 87(1/2) : 75-89. ——, 1959b. — Studies on New Zealand Elasmobranchii. Part IX — Scymnodon plunketz (Waite, 1910), an abundant deep-water shark of New Zealand waters. Trans. R. Soc. N.Z., 87 (3/4) : 271-282. ——, 1960. — Studies on New Zealand Elasmobranchii. Part XII — The species of Squalus from New Zealand and Australia; and a general account and key to the New Zealand Squaloidea. Trans. R. Soc. N.Z., 88 (3) : 519-557. ——, and PauL, L. J., 197la. — Heptranchias dakini Whitley, 1931, a synonym of H. perlo (Bonaterre, 1788) , the sharpsnouted or perlon shark, with notes on sexual dimorphism in this species. Zoology Publs Vict. Univ. Wellington, (54) : 1-14. ——, 1971b. — Cirrhigaleus barbifer (fam. Squalidae), a little known Japanese shark from New Zealand waters. Zoology Publs Vict. Univ. Wellington, (55) : 1-13. Lyncu, D. D., 1964. — First Australian record of Hexanchus griseus (Bonnaterre) 1780. The six-gilled shark. Mem. Nat. Mus. Vict., 26: 259-261. PHILLIPPS, W.J., 1946. — Sharks of New Zealand. Rec. Dominion Mus., 1 (2) : 5-20. PincHuK, V.I., and PeRMiTIN, YE. YE., 1970. — New data on dogfish sharks of the family Squalidae in the south-eastern Atlantic. J. Ichthyol, 10 (3) : 273-276. ROEDEL, P. M., and Riptey, W. E., 1950. — California sharks and rays. Fish. Bull. Calif., (75) : 1-88. SmiTH, J. L. B., 1967. — The lizard shark, Chlamydoselachus anguineus Garman, in South Africa. Occ. Pap. Dep. Ichthyol. Rhodes Univ., (10) : 105-114. STEAD, D. G., 1907. — In “Notes and Exhibits”. Proc. Linn. Soc. N.S. Wales, 32: 554. ———, 1963. — Sharks and rays of Australian seas. Sydney: Angus & Robertson. WHITLEY, G. P., 1940. — The fishes of Australia, part 1. The sharks, rays, devil fish, and other primitive fishes of Australia and New Zealand. Sydney: Royal Zoological Society of New South Wales. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 A Key to the Australian Genera of the Agaricales A. E. WOOD Woop, A. E. A key to the Australian genera of the Agaricales. Proc. Linn. Soc. N.S. W. 103 (4), (1978) 1979: 255-273. A key is provided to the genera of Agaricales found in Australia. The limits of the genera are those outlined by Singer, and most of the diagnostic characters have been incorporated in the key. A. E. Wood, School of Botany, University of New South Wales, P. O. Box 1, Kensington, Australia 2033; manuscript recerved 16 May 1978, accepted 21 February 1979. INTRODUCTION In recent years there have been great developments in the concepts of genera in the Agaricales. However, there has been little work carried out to apply these new concepts to the local agaric flora. This paper is an attempt to do this. The generic concepts used have been those of Singer (1975). In a few cases the generic limits are wider than those accepted by Singer. In almost all of these cases, the wider sense follows the use of British Check List (Dennis, Orton and Hora, 1960). The genera which have been included in other genera are as follows: Anellaria is included in Panaeolus Armillarzella is included in Armillaria Copelandza is included in Panaeolus Galerella is included in Conocybe Gerronema is included in Mycena and Omphalina Tectella is included in Panellus. Lentznellus and Schizophyllum are still included, despite their exclusion from the order by Singer. This conclusion is not universally accepted and for convenience they are still included in the key. In recent years I have collected and studied the local species and compared them with the genera accepted by Singer. In no case so far has it been necessary to propose a new genus. The genera included in the key are those that have been found in Australia together with those genera that may possibly occur here but have not so far been recorded. In this latter category are included large cosmopolitan genera and other genera where the distribution almost certainly includes Australia. Most of the collecting has been carried out in New South Wales, but there is reason to believe that the key will prove satisfactory for most of Australia. The key is based on the concepts outlined in Singer (1975) and Dennis, Orton and Hora (1960) and is dichotomous, with several genera being keyed out several times. Spore colours unless otherwise stated, refer to the colour of a spore print. Where possible, macroscopic features have been used, to allow the key to be used in the field. KEY ave ERMC DO Cy lalmMell aber omer AP Conn MORN role aces Soh (Care Seal (Cala aul aieen 2 lb. Fruit body soft, poroid (tough to woody SPEClesmSCEPROyMOIaCEAC) Means Weyer na | sho. e ie rc rei ce ates a egal ote 147 2a. (1) Lamellae narrow, fragile or flexible, crowded (space between lamellae < four times thickness of lamella) ; spore colour Nala OL GR eM ema Ea tate WAR RM 0 Fe <= raalnn Mend west ar aAaurs gat ya aj-wiathbaC 9 Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 256 A KEY TO THE AGARICALES 2b. Lamellae very thick, waxy, distant (space between lamellae > four times thickness ot lamella)sspoxe colour mostly whitey.) sai) ie ee eee 3 3a. (2) Spores brown, smooth, elongate; cystidia present, usually large and encrusted; basidia not very elongate (L/d<6) Gomphidius 3b. Spores white, smooth or rough, non- amyloid:);basidiaelongateon NOt ile. eee eee ee 4 4a. (3) Basidia not very elongate (£/d<6) ; spores: smooth: on TOUR eke ini in ON Mer aa oN re arc usa 4b. Basidia elongate’ (€/d>6)); spores smooth) 3022 te: 6 5a. (4) Spores elongate, smooth; cap margin grooved; pigment present that turns green in alkali Anthracophyllum 5b. Spores globose, spiny; cap margin smooth; no unusual pigment present Laccarva 6a. (4) Lamellae shallow, decurrent, anastomosing or reduced Cantharellus* (Cantharellaceae) OU 6b. Lamellae well developed, decurrent or NOt) NOtianastOmMosingy, ai) Uke! es tee I al eee eee Z 7a. (6,42) Lamellar trama distinctly bilateral; cap viscid; lamellae usually decurrent; stipe apex usually rough with dots; veil often present Hygrophorus Wb veamellarstramacnot bilateral) (i oe sae) ope ee en oa ieee 8 8a. (7) Lamellar trama strongly interwoven ; lamellae often decurrent; fruit body not bright red or bright yellow; cap never viscid; stipe smooth Camaro phyllus 8b. Lamellar trama regular, mostly of wide hyphae; lamellae decurrent or not; colour of fruit body usually bright red or bright yellow; cap often viscid; stipe smooth Hygrocybe 9a. (2) Flesh soft, fibrous; spores of various colours, amyloid, dextrinoid or non- AMMVLOL ome ii i ete oe aaantnt a urine: Oia bik scala Nc serie gc NNR ame 11 9b. Flesh granular (containing sphaerocysts) ; spores white to pale buff, amyloid, OLMAMENted ey ee On ly ee ka he a eau aan ais onlis obs iicg ee ei s 10 10a. (9) Cap with latex when broken; intermediate lamellae always present Lactarius 10b. Latex absent; intermediate lamellae mostly absent Russula Placa (O)eStipercentralty a) oan duno ie eae hop ne nea ci cae ae ate 28 Libs wStipeseccentrics lateralior absemty: Wimai ne cays cer cee ed ieee 12 * This is not a true agaric, but it is included because of superficial similarity which might cause it to be confused with this group. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 12a. bs 13a. 13b. 14a. 14b. 15a. 15b. 16a. 16b. 17a. 17b. 18a. 18b. 19a. 19b. 20a. 20b. 2la. 21b. 22a. 22b. A. E. WOOD (11) Spores mainly white or pale brown; almost always on wood Spores pink or purple; (12) Spores purple; cuticle filamentous, gelatinized ; on wood Spores pink and ezther polygonal or with ridges (12) Fruit body soft and fleshy Fruit body tough and leathery (14) Spores pinkish-brown to ochre- brown Spores white, non-amyloid (15) Spores pinkish-brown to clay- brown; lamellae not anastomosing ; trama ><* regular Spores cigar-brown to ochre-brown ; lamellae decurrent >< anastomosing, readily separated from cap flesh; trama bilateral; sometimes on the ground (15) Gelatinized layer in flesh; encrusted, thick-walled marginal cystidia present No gelatinized layer in flesh; + cystidia of various types (17) Cap surface with stiff dextrinoid hairs; thin or thick-walled cystidia present; fruit body sessile, with constricted base Cap surface of unspecialized hyphae (18) Fruit body resupinate; thick-walled cystidia absent Fruit body sessile or stipe eccentric (19) Lamellae distant (space between lamellae > four times thickness of lamella) ; cap margin grooved; spores cylindric; cystidia absent; pigment present that turns green in alkali Lamellae crowded (space between lamellae < four times thickness of lamella) ; cap smooth (20) Spores ellipsoid to cylindric; stipe eccentric, rarely absent; >< thick-walled cystidia present Spores sub-globose (21) Fruit body sessile; thin-walled cystidia present or absent; cap thin Stipe eccentric; cystidia absent; cap fleshy, luminescent Melanotus Co Crepidotus Paxillus Hohenbuehelia Chaetocalathus CC SO ST Resupinatus oe 8 © © © © © ew 8 ee wt ww ww kl Anthracophyllum Ce Pleurotus Nothopanus Omphalotus * >< = more or less. Proc. Linn. Soc. N.S.W., 103 (4) , (1978) 1979 258 A KEY TO THE AGARICALES 23a. (14) Lamellae splitting longitudinally Schizophyllum 23be 7 amellacmmot;splitting oe Se Oite Mle Peck aecatens) cuetaue tame eat etnies Z4a. ¢.(23)) Mangin) ofvlamellae entire! panera) Ni. wea Rein yl eters deat 24b.5-“Maroinvot lamellae :seriates mayan fine eee ae era ee ani ia 25a. (24) Spores amyloid; cystidia absent Panellus Z5b2) Spores mon army lOidye ap: oy eee ite sa tant as en ee 00 Liat eens gece eaten cope 26a. (25) Cystidia present; lamellae crowded Panus 26b. Cystidia absent; lamellae distant; cap margin grooved Anthracophyllum 27a. (24) Spores non-amyloid, cystidia absent Lentznus 27b. Spores amyloid, cystidia present Lentinellus 28a. (11) Spores white or pale green or pale jOSDOU aU a Seta LL CLA MON MMS ALC BRM an Aaa eae eae ea IGE 28b.)4), Spores deepipin kis ae cue anes. eRe ei UO EEE, 28c. Spores rust, clay or cigar brown (not chocolate) UR SIE AY MMR eae da icp a EAC OM OR eam ee uaa a ht i 28d. Spores chocolate, cocoa brown, purple or |B) EXO) ye eRe recs aed aan Mi ie OPN oat a Unione CMA MEIER CURIA a Nic Atuly 29a. (28) Stipe with ezther volva, volval remains or bulbous base and/or a distinct annulus!(at leastiin) young) Specimens) ye ne ea eee 29 ban Stipeswith neztenwolvasmor ammMulusy rae a to. iene ae 30a. (29) Stipe with ezther volva, volval remains or bulbous base; annulus mostly present; spores amyloid or not, never dextrinoid Amanita 30b. Stipe with no volval remains, but with a distinct sammuluisy sie a OO Oa Oh cra len) Seal lie, ae eee Silat’. (30); lamellae adnatexto decurment iyi apts We eae ve eee 31b. Lamellae free; fruit body normally on GIVE VO TOMI Hiya eal geal cea ahh UNG aA eGo care Reed eats 2 ea 32a. (31) Cap cuticle filamentous; fruit body normally onowoOd: i ysRi heey iar Wea e Piet aris Maral MNEs Wasser tsa ae neta rig: 32b. Cap cuticle cellular or with a covering of cellular granules; fruit body mostly on (oy eMiea Rey Vo ELE caer anna egive Aenea MCC CPIM Sunt CL Mden tanner! Mice, Che in eed 33a. (32) Cap smooth or with small scales Armillaria 33b. Cap with prominent, large, erect, conical warts Cyptotrama 34a. (32) Cap mostly viscid; cap cuticle cellular; large cystidia present Oudemansiella 34b. Cap not viscid; cap-and stipe with a covering of spherical cells in granules; cystidia not prominent Cystoderma 35a. (31) Cap viscid; lamellar trama bilateral; spores small, sub-globose, non- amyloid Limacella 35b. Cap dry; lamellar trama never bilateral; Spores dextrimord aia ieee Mii) iy aU nes er ee cee Sarin ete lee 36a. (35) Lamellae and spores >< green Chlorophyllum SOD ce, SPOTES WHIGE es A Mai NAN Ga EN ea a a MN dae Nc rai aie ey Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 37a. 37b. 45b. 46a. 46b. A7a. 47b. 48a. A. E. WOOD (36,53) Spores metachromatic in cresyl blue, with prominent germ pore; cap fleshysorsim eM bia OUSIe wang hh oi Mis dite de Shane Ce eaves oan Spores not metachromatic in cresyl blue, without prominent germ pore; cap fleshy ................. (37) Disc of cap with palisade structure; clamp connections present; cap large, fleshy; spores without metachromatic plug Macrolepiota Disc of cap without palisade structure; claimprconnections absenty on Fe ran Wal hares Mi etiu Shail (38) Cap large, fleshy; spores without metachromatic plug Leucoagaricus Cap membranous; spores with metachromatic plug Leucocoprinus (37) Cuticle cellular but not a palisade; spores with thin walls; clamp connections present or absent; cystidia usually present Lepzota Cuticle flat filamentous; spore walls somewhat thickened; clamp connections absent; cystidia absent Pseudobaeospora (9) slcamellace>—<— strongly decurrent <7 993) fel s ameliae mot istrone ly Gecurnremte rye ema. io Piet Sei eis 3) (41) Lamellae thick; basidia very long (LA COROT) SG aN orate eg ee eer ier 2 eS a ceo ae Lamellae thin; basidia not unusually OIC eerie aie tometer yee cel A SAAS Say es ie cee oes Al G2))-Spores non-amyloid or dextrinoidi) (0)... i) so SPOLESAATVOLC nee Aes ol la aah Rear ae iescss Sylar ae (ES) ESHORES SIMNOOC ane Win Marini faunas eld eel Wise aise i etal ia es Spore wall rough, uneven or INETSKOREMEOUS Iai cyte ean (tos un meh ena li eee auRlans Cuma Sables hore (44) Small species (cap diameter usually < 2 cm); stipe often tough to cartilaginous; lamellar trama irregular; clamp connections present or absent Omphalina Large species (cap diameter usually > 2 cm) stipe typically fleshy; lamellar trama regular or irregular; clamp connections [SESE Ewart Rey sh) cain hs coe Me Biae cic e aia Nels ey GaN « (45) Lamellae repeatedly forked Hygrophoropsis mamelideymot forked ay. ih ox Murcia tean hay esate WDC) cscs yelehac edu (46) Fruit body bright yellow to orange, luminescent; lamellar trama irregular to >< regular; on wood Omphalotus Fruit body not bright coloured, not luminescent; lamellar trama >< regular; mostly on the ground Clitocybe (44) Spores with heterogeneous or uneven walls; cystidia present; fruit body drab coloured Fayodia Proc. Linn. Soc. N.S.W., 103 (4) 259 , (1978) 1979 260 A8b. 49a. 49b. 50a. 50b. 5la. 51b. 52a. 52b. 53a. 53b. 54a. 54b. 55a. 55b. 56a. 56b. 57a. 57b. 58a. 58b. A KEY TO THE AGARICALES Spores distinctly spiny; cystidia absent ....... PR ae eae ee 49 (48) Spores white to cream Clitocybe Spores pink Lepista (43) Stipe tough, with yellow to rust basal mycelium; fruit body somewhat reviving; spores smooth, thin-walled; cystidia present; on wood or humus Xeromphalina Stipe more fleshy; basal mycelium not coloured; fruit body not reviving; cystidiapresent ior absent, ah hes Wii oR anes SES Riey carbines nea. 5] (50) Cystidia absent; spores short- ellipsoid to sub-globose, smooth, thin- walled; on wood Clitocybula Cystidia present; on wood or on the GROUING hE CC ale iho b Bays ns sale Gamat Bie oi Minet Rceyae eR NEA 52 (51) Spores with smooth, simple wall, ellipsoid to short-cylindric; on wood Clitocybula Spores with thick, more or less uneven, wall, >< subglobose; on wood or on the ground; cuticular hyphae sometimes gelatinized Fayodia (41); Lamellae free;spores dextrinoid), 7 2223 a ae 37 Lamellae free, adnate or sinuate; spores not dextrinoid (rarely dextrinoid and then lamellae not free) (53) Cap and stipe with a covering of granules consisting of spherical cells; spores amyloid, non-amyloid or dextrinoid Cystoderma Cap and) stipe without axcellular covenme) 4 5 oe Sl 55 (54) Lamellae adnate; spores thick walled, dextrinoid, without germ pore; stipe fleshy, without annulus; cap fleshy, smooth, viscid; cuticle filamentous Hebelomina Spores not dextrinoid; lamellae free, adnate or sinuate (55) Spores rough (or at least heterogeneous) sii) aie ete) EAN CONS ARN aA a entero ea bee 57 Spores smooth (56) Spores amyloid Spores non-amiyloid yey, Pei ae a deere Waele airay th crieed Aeshna er 59 (57) Hyphae without clamp connections; large pointed marginal cystidia, with crystal coated apex present (may be rare) ; cuticle filamentous; spores with plage; fruit body usually fleshy with sinuate lamellae Melanoleuca Hyphae with clamp connections; prominent cystidia absent; cuticle filamentous; spores without plage; fruit body usually fleshy with sinuate lamellae Leucopaxillus Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 59a. 59b. 60a. 60b. 6la. 61b. 62a. 62b. 63a. 63b. 64a. 64b. 65a. 65b. 66a. 66b. 67a. 67b. 68a. 68b. 69a. 69b. 70a. A. E.WOOD (57) Lamellae thick, distant, (space between lamellae > four times thickness of lamella), broadly adnate; cuticle filamentous; spores without plage; basidia without carminophile granules Lamellae thin, crowded (space between lamellae < four times thickness of lamella), often sinuate; spores without plage (59) Cuticle cellular, usually above a gelatinous layer; large prominent cystidia present; lamellae attached; clamp connections mostly present Cuticle filamentous; cystidia absent or present (and then not very large) ; clamp connections present (60) Basidia without carminophile granules Basidia with carminophile granules; spores white (61) Spores pale pink; spore wall thin, coarsely to finely roughened Spore print white; spore wall thick, heterogeneous or uneven (61) Fruit body grey Fruit body bright coloured (rarely white) (56) Spores amyloid Spores non-amyloid (64) Cystidia rare or absent Cystidia present, conspicuous (65) Fruit body large (cap diameter usually > 2 cm), fleshy, on the ground; lamellae often sinuate Fruit body small (cap diameter usually < 2 cm), on wood (66) Hyphae with clamp connections Hyphae without clamp connections (66) Lamellae well developed Lamellae mostly reduced to veins; fruit body white, fragile (65) Fruit body with large pointed marginal cystidia with crystal coated apex (may be rare) ; hyphae without clamp connections; fruit body mostly large, fleshy; lamellae usually sinuate Fruit body with conspicuous or inconspicuous cystidia, but these never crystal coated; hyphae with clamp connections (69) Cystidia not conspicuous; fruit body mostly large (cap diameter usually > 2 cm), fleshy; lamellae often sinuate Laccaria Oudemansiella Lepista Fayodia Lyophyllum Calocybe a Cn Cn ec Leucopaxillus Melanoleuca Clitocybula Delicatula Melanoleuca Od a°0 Db OloG Oo G6 Oo Oo Oo DO 5 Leucopaxillus Proc. Linn. Soc. N.S.W., 103 (4) 261 , (1978) 1979 262 70b. 71a. Tilo 72a. 72b. gas 73b. 74a. 74b. 75a. 75b. 76a. 76b. 77a. 77b. 78a. 78b. 79a. 79b. 80a. 80b. 8la. 8lb. 82a. 82b. A KEY TO THE AGARICALES Conspicuous cystidia present; fruit body mainly.smali (cap diameter usually < 2 cm), cap conical; lamellae ascending, >< free (64) Cap cuticle normal filamentous, without hairs or cystidia Cap cuticle diverticulate to cellular, or filamentous with hairs or cystidia (71) Stipe fleshy; fruit body mostly large (cap diameter usually > 2 cm) ; lamellae typically sinuate Stipe tough; fruit body seldom large; lamellae typically adnate (72) Cystidia present, marginal, >< globose; on wood Cystidia inconspicuous or absent; usually on the ground (73) Baidia without carminophile granules; cystidia absent Basidia with carminophile granules (74, 77) Fruit body grey (rarely white, and then basidia long) Fruit body bright coloured (when white, basidia not long) (72) Gloeocystidia present Gloeocystidia absent (76) Basidia without carminophile granules Basidia with carminophile granules (77) Stipe not insititious; cuticle not gelatinized; fruit body not reviving Stipe insititious; cuticle gelatinized or not; fruit body reviving (78) Cuticle gelatinized or stipe with black rhizomorphs; odour usually unpleasant Cuticle usually not gelatinized ; rhizomorphs absent; odour not unpleasant (71) Cap cuticle filamentous Cap cuticle diverticulate to cellular (80) Cap with long stiff hairs; fruit body fragile, mostly white; cap dry Cap with cystidia (81) Cap with large cystidia (length up to 60 um) ; fruit body large (cap diameter usually > 2 cm), coloured; on wood; cap viscid Cap with small cystidia (length up to 30 um); fruit body small (cap diameter usually < 2 cm) delicate, white; cap >< hygrophanous Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 Mycena SSCS R ORE TTY PPR oRRGI RTOS a 72 Bae AU raid St RSLs aM te 80 SE ni A ret ee ef a3 ERROR MRC Rohe yarn 76 Tricholomopsis 2A INSU taMRe ta Sepa man he ANB 74 Tricholoma Ly ai Sie ane arc Re oleae eM Se 75 Lyophyllum Calocybe Lactocollybia TULARE UN a ose ORE as 77 Ae Neate rie hier gh 78 sel SORIA ich Gnaiterm an RMR M 75 Collybia cal egg Gc SE Ree Jaen a 79 Micromphale Marasmiellus 1 SLR DOR Te OE NMER s Chapa 81 Alek Nts SPAIN, BNE Resa ese ccs 83 Crintpelles Te Cerise) Len neara es Citra here 8 nee 82 Flammulina Hemimycena 83a. 83b. 84a. 84b. 85a. 85b. 86a. 86b. 87a. 87b. 88a. 88b. 89a. 89b. 90a. 90b. Ola. 91b. 92a. 92b. 93a. 93b. A. E. WOOD (80) Fruit body small (cap diameter usually < 2 cm), white, non-reviving; lamellae with no marginal cystidia; cap cuticle diverticulate Fruit body not combining these characters (83) Cap cuticle cellular to sub-cellular Cap cuticle nodulose to diverticulate (84) Cystidia inconspicuous or absent; basidia with carminophile granules; fruit body not reviving Conspicuous cystidia presént; basidia without carminophile granules (85) Fruit body not reviving; stipe fleshy; sub-cuticular layer of cap usually gelatinous Fruit body reviving or not; stipe tough to cartilaginous; sub-cuticular layer of cap not gelatinous (86) Fruit body reviving; cap cuticle without cystidia; cells of cuticle smooth or rough to irregular; clamp-connections mostly present Fruit body not reviving; cap cuticle with cystidia; cells of cuticle regular, smooth; clamp-connections absent (84) Stipe not insititious; fruit body not reviving; marginal cystidia present or absent, not conspicuous Stipe insititious; fruit body usually reviving; marginal cystidia mostly present (88) Stipe tough but not cartilaginous; cap cuticle diverticulate but not rough Stipe thin, cartilaginous; cap cuticle diverticulate and roughened (13, 28) Stipe eccentric, lateral or absent and spores ribbed Stipe usually central (90) Spores polygonal, smooth; lamellae not free; cap and stipe not separable; mostly on soil Spores smooth or rough, not polygonal (if somewhat angular then also rough) (91) Spores rough; lamellae not free; cap and stipe not separable Spores smooth (92) Clamp connections absent; cystidia present or absent; cuticle >< parallel filamentous; spores ovoid to somewhat angular Clamp connections present; cystidia absent; cuticle interwoven filamentous; spores ovoid Hemimycena Oudemansiella i i i Marasmius Pseudohiatula Collybza Marasmiellus Marasmtus Clito pilus eo 8 © © © 8 we ew ew 8 ew ew wll ee ote iol olive tet e) GelielLeure Melie) elke) (el Ke Rhodocybe Lepista Proc. Linn. Soc. N.S.W., 103 (4) , (1978) 1979 264 A KEY TO THE AGARICALES 94a. (92) Large, elongate, thin-walled, hyaline cystidia on lamellae and cap surface (length up to 60 ym); lamellae sub-free; cap and stipe not easily separable Macrocystzdia 94b. Large cystidia absent; lamellae free; cap andistipe easily sepanables iy igi pcajone Uieusie ey Co Ne AMV ian ene tir. 95 95a. (94) Well developed cup shaped volva present; onthe ground: or woodin. 5 ii Mier es Ciena ete 96 95b. Volva absent; lamellar trama inverse; Spores mon-ammylordiy ate ein iy sage! Ne ACG Boer teria) aero seein 97 96a. (95) Lamellar trama inverse; spores non- amyloid; on the ground or wood Volvariella 96b. Lamellar trama bilateral; spores amyloid; on the ground Amanita 97a. (95) Annulus present Chamaeota 97b. Annulus absent Pluteus 98a. (28, 114) Fruit body fragile, soon decaying; cap viscid, striate or pleated; cuticle cellular; cystidia not abruptly capitate; lamellar trama regular; stipe with no brown pigment towards base; spores rust-brown, occasionally dull- brown Bolbitzus S8be: Bruit body not delicate. frapilei in sin aches eM ey ere eens epee 99 99a. (98) Stipe with membranous ring at TIVABUITLE Yai UN an Say cS ys ile FAO dS NL ea cei ie ARH leg neg 100 99b. Stipe with cobweb partial veil or veil absent (cobweb partial veil, when present, mostly only visible in young stages, adult specimens may show fine fibrillar remains on) stipe) eee) view week ere eet) es Skene ieee Benen 108 1004.7: (99) Cuticle filamentous.e: sy ‘Pisces iii Sanaa naan wee eit hr pare 101 hOObDEs “@uticle cellular ya) Use a ea ogee ean eens eb ueteaa gs ene Ege 106 1Ola 700) Sporessmoothiac yay ee ines ie 1 Me (BP cnet ea ak aoe eats 102 LODE SC SPOKES rOU eda i ee Oo Ii ain cama ah Dips Ntaalao emul Oops Ueda eS 105 102a. (101) Lamellae thick, distant (space between lamellae > four times thickness of lamella), decurrent, often waxy; spores elongate, without germ pore; cystidia present, usually large and encrusted (length up to 60 ym) ; cap mostly viscid Gomphidius 102b. Lamellae thin, crowded (space between lamellae < four times thickness of lamella) eenotwaxypmrarelyadecurnent, vai) ie fer eins oe eeu nery aaa 103 103a. (102) Spores truncate, with prominent germ pore; chrysocystidia absent; cap hygrophanous, not scaly; mainly on wood Kuehneromyces 103b. Spores without prominent germ pore; chrysocystidiar present or absent, ) 7) sale ayn iene eels an ieee 104 Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 104a. 104b. 105a. 105b. 106a. 106b. 107a. 107b. 108a. 108b. 109a. 109b. 110a. 110b. llla. 111b. 1]2a. 112b. A. E. WOOD (103) Mainly on wood; stipe mostly >< scaly; lamellae not decurrent; spores ovoid or non-ovoid; chrysocystidia often present; cap not hygrophanous; fruit body usually large, fleshy (cap diameter usually > 3 cm.) Pholiota Mainly on the ground; cap or stipe not scaly; chrysocystidia absent; spores ovoid; fruit body mostly small (cap diameter usually < 3 cm) Galerina (101) Fruit body large (cap diameter usually > 3 cm), fleshy; spores without plage; veil ezther double, forming annulus on upper part of stipe and annulus or belts on lower part of stipe or single Rozites Fruit body mostly small (cap diameter usually < 3 cm), fragile, striate; spores with plage; veil single Galerina (100) Spores rust-brown, smooth, with germ pore; cystidia not abruptly capitate Pholzotzna Spores buff-brown; cystidia not abruptly CAP ILACC NE rupee bel neaulihe) A edt cika! Vel Citivas yucca da thang ne ON cr (106) Spores smooth, with germ pore Agrocybe Spores rough, without germ pore, without plage Descolea (99) Cuticle cellular or sub-cellular or with abundant cystidia; mostly small species (cap diameter usually < 3 cm) ; Spokes SIMOOtM Or LOUgIy: hen daerivea adel = ine veueera tee ert Mal na / ae Cuticle filamentous, without cystidia; small or large species; spores smooth or MOUS y er etaenev paste g 8s Salon, Wiecaiihd sige) VON gS Nad fetbaikan aaa de sae he (108) Spores rust-brown to cigar-brown.................. Sponesychocolate-browm ney Gi ale agile) aie cok ey ye ea ie (109) Cuticle strictly cellular, with or without cystidia; spores smooth, with prominent germ pore; cystidia >< ARUP tyACa ICAL Gamay hh Mibkna Rr a y unetOEN IS. MaMa ins CK ka, Cuticle sub-cellular, with or without cystidia; spores smooth or rough, without prominent germ pore; cystidia not AD KUIP ClyCapibate i evi iid jbl es aneemey Ace e MANNS) Ses coh Pada elcep as ols (110) Spores cigar-brown; cystidia not abruptly capitate; cap without cystidia, convex to flattened Agrocybe SPORESHEUStDROWMy anyon vrai Pieaigiicel we anel luewspisiecthaliene’ Shee) swede) siete (111) Cystidia abruptly capitate; stipe brown towards the base; cap without cystidia Conocybe Gystidianot abruptly scapitat cosmo ve legen inca ten sesievel err Proc. Linn. Soc. N.S.W., 103 (4) , (1978) 1979 266 113a. 113b. 11]4a. 114b. 115a. 115b: 116a. 116b. 117a. 117b. 118a. 118b. 119a. 119b. 120a. 120b. 12la. 121b. 122a. 122b. 123a. A KEY TO THE AGARICALES (112) Stipe brown towards the base; cystidia often present on cap Pholiotina Either stipe white to the base or cuticle not, strictly; cellubans 072301 Uae pa cae Ra etc iy tyae oe SSO ancre ye 114 (113) Stipe white to the base; spores WICH PEM ADOLES fis 1 rsh pire Nia SUNG etn aaa coe ade a Al 98 Stipe coloured; germ pore inconspicuous Or absent ; ‘cystidialoften presentionicap ys en a ees oe 115 (110, 114) Spores rough, cigar-brown; cap cuticle >< a layer of cystidia or >< cellular Alnicola Spores smooth, cigar-brown or rust- brown (115, 124) Fruit body hygrophanous; spores cigar-brown; cap cuticle a layer of cystidia Simocybe Fruit body non-hygrophanous; spores rust-brown; cap cuticle of broad elongate cells, heavily encrusted, often in chains Phaeomarasmius COS) Sporesismoot hays: et a tie ra leet Se ee COG heen eat 118 Spores rough or nodulose (this may need tobe determined’ under oilhimmersion) 9) aes ee eye re 132 (117) Spores clay-brown or cigar-brown OF PLEY; PLOW Mie WO Se ree ea VRE IC NT ACRES HD ee pe aa 119 Spores: rust DrOwmnterye et rey ere MOLE ne Ses OAeeNct ts Rig ee crete ca ZO (118) Spores clay-brown or cigar-brown; chrysocystidia: seldom) presemean | sui: 0s se ye eis ree oan 124 Spores some shade of grey-brown; chrysocystidia-often present 21) ik) (yo enn cs neces at oa alte eh Atanas 141 (118, 131) Spores truncate, with prominent germ pore; chrysocystidia absent; fruit body hygrophanous; lamellae not free Kuehneromyces Spores not truncate, without prominent germ pore (narrow indistinct pore may be present in some cases) (120) Spores thin-walled, often collapsing in water, >< hyaline sub. muicr. Spores usually not collapsing in water, coloured sub. micr. (121) Clamp connections present ; lamellae broadly adnate to sub- decurrent; marginal cystidia narrow; facial cystidia absent Tubaria Clamp connections absent; lamellae ascending, >< free; cap conic to hemispherical, normally striate; marginal cystidia narrow; facial cystidia present Galerina (121) Fruit body mostly medium to large (cap diameter usually > 3 cm); cap >< Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 123b. 124a. 124b. 125a. 125b. 126a. 126b. 2ae 127b. 128a. 128b. 129a. 129b. 130a. 130b. A. E. WOOD convex; lamellae usually attached; ezther cap scaly and/or chrysocystidia present and/or spores non-ovoid Pholiota Fruit body mostly small (cap diameter usually < 3 cm); cap >< conical; lamellae ascending, >< free; cap not scaly; chrysocystidia absent Galerina (119) Cap cuticle filamentous, of the TNO Ten cll (3h fC eta wna OR ee 0 NN nse sh oecr R e ole I SIE co ae Cap cuticle of >< barrel-shaped cells or Chraimsrotbroaduelongatencellliss gi, | 9) yes la ia ae (124) Lamellae decurrent; lamellar Biabinag il abelallaetewe wets a eee ema tA ees Naso ra Niey 2 4 Lamellae rarely decurrent; lamellar Balance Cull alii was ty MieeRIN tees, Miwa Tea erento hs sean (125) Lamellae thick, distant (space between lamellae > four times thickness of lamella), often waxy; spores elongate without germ pore; cystidia present, usually large and encrusted; cap mostly viscid Gomphidius Larmellae thin, crowded (space between lamellae < four times thickness of lamella), not waxy, >< anastomosing; CAREC aVISCIMat am tea | NAN Rigen serena acy Romie wee cets (126) Lamellae readily separating from flesh; cystidia often present, not encrusted; spores not elongate Paxillus Lamellae not readily separating from flesh; cystidia present, often encrusted ; spores elongate; cap cuticle turns blue with ammonia Phylloporus (125) Cap mostly conical, generally radially fibrillose or splitting; cystidia striking, ezther thick-walled and with crystal coated apex or thin-walled, naked, cylindric; spores ovoid, without germ pore Inocybe Cap mostly convex, not radially ‘ilorillowes jystichen moe sun my — ga be ae ecb osae (128, 141) Spores not truncate and without prominent germ pore Spores truncate or with red line at junction of wall layers when mounted in KOH (129) Spores thin-walled, often collapsing in water, >< hyaline sub. micr. Tubaria Spores firm-walled, not collapsing, coloured sub. micr., without reddish line at junction of wall layers when mounted Proc. Linn. Soc. N.S.W., 103 (4) 267 , (1978) 1979 136b. 137a. 137b. 138a. 138b. 139a. 139b. 140a. A KEY TO THE AGARICALES in KOH; ezther cap scaly and/or chrysocystidia present and/or spores non- ovoid (129) Chrysocystidia absent; spores truncate, with prominent germ pore shade of grey-brown; wall layers brown or cigar-brown (132) Spore wall punctate due to embedded spines Spore wall with true spines or warts (133) Spores rust-brown Spores clay-brown or cigar-brown cobweblike; spores rough-warted clamp connections present; spores without plage but then clamp connections absent) pore; cystidia present; cap cuticle normal filamentous, without cystidia cm) dry; spores without germ pore; of cystidia or >< cellular lamellae free; spores cocoa-brown to purple-brown; annulus present Fruit body otherwise Pholiota Chrysocystidia present; spore print some spores mounted in KOH show reddish line at junction of (117) Spores angular, nodulose, clay- Inocybe Spores spiny or wartedsebut: not moduloseye aise 5 rien eee eee 133 Tubaria 5 ialconeh tos CERES CAN 134 1 ESSA NE ee ats ae iy aE 135 Mere SM ea nar td aI Ae 137 (134) Cystidia typically absent (when present, >< globose, thin-walled) ; veil Cortinarius Cystidia, typically, present, not globose’. 95) 5. 136 (135) Fruit body bright coloured, often on wood; cap >< convex; veil present or absent (always on wood when present) ; Gymnopilus Fruit body mostly not bright coloured ; cap conic to hemispherical, normally striate; lamellae ascending; mostly on the ground; clamp connections present; spores with plage (rarely without plage, Galerina (134) Cap often large (diameter usually > 3cm), viscid; spores without germ Hebeloma Cap usually small, (diameter usually < 3 cystidia present; cap cuticle >< a layer Alnicola (28) Cap and stipe readily separable; Agaricus (138) Fruit body normally autodigesting; lamellae parallel-sided; spores black or Coprinus rarely chocolate Fruit body not autodigesting (139) Cuticle filamentous; spores deep brown to violet; fruit body not very fragile ee 8 we ew ww ew ww wt lw ee Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 Ce 140b. 14la. 141b. 142a. 142b. 143a. A. E. WOOD Cuticle cellular; spores black or deep bowtie DOG sina Giles Ge limit pokey tare ch ak 4 il eur a at (140, 109, 119, 131) Spores drab-brown to blackish-violet, often non-ovoid; spores sub. micr. in KOH greyish or with a reddish line at the junction of the wall ESPEIRS Ta NR EURS CRIS pa hea Ne Ri eer i Oey nd ea a Spores yellow-brown to deep rust-brown or grey rust-brown, rarely non-ovoid; spores sub. micr. in KOH yellow-brown, without a reddish line at the junction of Blvegweallibliay Cis beara Sade Ab er eet paolo rhea Nes ene ual aah a (141) Sub-cuticular layer of cap sub- cellular; cap typically dry, with no separable pellicle; chrysocystidia always present; often tufted and on wood; veil cobweb-like, rarely annulate; lamellar trama regular, of narrow hyphae (rarely also with a central strand of swolllen cells) Naematoloma Sub-cuticular layer of cap not sub- cellular; cuticle typically a separable pellicles chnysocystidialjpresent or/absent)) +. 95...4.5..24.-% (142) Chrysocystidia normally present (where absent, stipe with glutinous covering) ; annulus present (may be fugacious) ; lamellar trama of interwoven hyphae of unequal diameter Stropharia Chrysocystidia absent; veil mostly sparse or absent; lamellar trama sub-regular Psilocybe (140) Spores discolouring to a grey colour (swb. mzcr.) in conc. H,SO,; lamellae not mottled; spores smooth (srenreelhyy TROON 1) eS eg tne ty en Mey Ae A Ble oR oy o, Spores not discolouring in conc. H,SO, (sub. micr.) ; lamellae mottled; spores SIMGOEMOTOUCIG ik. eal: peared Manteca “Yau he ne Ose Sis vane saree. (144) Cap smooth; hymenium without pseudoparaphyses; stem white Psathyrella Cap pleated, grooved; hymenium with pseudoparaphyses Coprinus (144) Spores smooth Panaeolus Spores rough Panaeolina ()eSporesismooth white orcolouned’= fen) sees SoOress owen, lola, wre imeendhy GO 9 eee bebo cs oe eorex QUES POKeSiWiite cs aie Stance aM cam ull rits Cenk Aeron ERS A Spores yellow, pink, fawn, brown, olivaceousiornblackauey tu. We Mie remade yk Abe eR ey fee eo (148) Fruit body fragile, white; spores ellipsoid to sub-globose, amyloid Filo boletus Proc. LINN. Soc. N.S.W., 103 (4) , (1978) 1979 270 A KEY TO THE AGARICALES 149b. Fruit body fleshy to somewhat tough; surface of cap buff to brown; spores cylindric, non-amyloid Polyporus arcularius* (Polyporaceae) 150a:, (148). Sporesinotblacky i caval ie hs ae Une ore Aone eee Mie Nel 150b. Spores black and ezther very large or with a very thick wall PAR GBN I~ scoters he anlar et chute SARIN cee aes eR Mie 159 15la. (150) Clamp connections present and easily found(spores short’) ei gy 0 Wel cis penne ccs nee Asin aay ote x2 151b. Clamp connections absent, sparse or present (when present, spores elongate and tubesyarranged radially iim Ay il) hale sire en ere on rea 153 152a. (151) Tubes >< free around the apex of the stipe; spores yellow Gyro porus 152b. Tubes >< decurrent; spores brown to olive-brown Gyrodon 153a. (151) Evzther pores arranged radially or fruit body entirely red to pink Suzllus 153b. Fruit body with none of the above Characters ee mei em eN kil as il VN ee es ol bau ees eae Sine ese arg nea an 154 154a. (153) Spores ferruginous-brown, fawn or pink; flesh never blueing Tylo pilus 154b4 Sporesyolivaceous- brown | 1G) yeinatae hs tl | ae gap ae ee ee ether Dele a 155 155a. (154) Stipe scabrous from squamules which are white at first, but soon discolour black or brown; stipe rarely cylindric or ventricose; pores never red Leccinum E5Sb.7¢ Stipe motscabmousis (lie. i), aoe ies) PEIN ati eneatra ty anew pt cme 156 156a. (155) Veil pulverulent to arachnoid, sulphur yellow Pulvero boletus 156b 2: WVeilvothenwise ite) 1) ns0 tes iA gesbede ol, FSET atlaes Res cis aa csi teal ofa aetna tea a WSs 7/ 157a. (156) Cap viscid; cuticle not cellular Pulvero boletus 157b. Cap viscid or dry, when viscid, cuticle Cel ag leg aes aR. a pa igky co Sa OA os Ie 158 158a. (157) Stipe reticulate and/or tubes red at first; tube trama with strongly divergent hyphae which are much paler than the central tissue Boletus 158b. Stipe not as above; tube trama with slightly divergent hyphae which are only a little paler than the central tissue Xerocomus 159a. (147, 150) Spores globose to sub- globose; cap woolly or warted; spores almost black Strobilomyces L59b.; “Spores: Clom gate wigs 5 i) eka epee aa oe aS SO Mica 160 160a. (159) Spores with embedded spines or pegs (occasionally smooth) ; young tubes white to pale grey; spores red-brown to dark-brown Porphyrellus * This is not a true agaric, but it is included because of superficial similarity which might cause it to be confused with this group. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 A. E. WOOD 271 160b. Spores with longitudinal ridges (occasionally with embedded spines or smooth) ; young tubes yellow; spores almost black Boletellus ALPHABETICAL LIST OF GENERA AGARICUS L. ex Fr., Syst. mycol. 1: 8. 1821. AGROCYBE Fayod in Ann. Scz. nat. (Bot.) VII9: 358. 1889. ALNICOLA Kuhner in Botanzste 17: 175. 1926. AMANITA Pers. ex Hook., Fl. scot. 2: 19. 1821. ANTHRACOPHYLLUM Ces. in Att? Accad. Sci. fis. mat. Napoli 8 (3) : 3. 1879. ARMILLARIA (Fr.) Kummer, Fiihr. Pilzk. 25, 134. 1871. BOLBITIUS Fr., Epicr. 253. 1838. BOLETELLUS Murr. in Mycologza 1: 9. 1909. BOLETUS Dill. ex Fr., Syst. mycol. 1: 385. 1821. CALOCYBE Kuhner in Bull. mens. Sac. linn. Lyon 7: 211. 1938. CAMAROPHYLLUS (Fr.) Kummer, Fuhr. Pilzk. 26, 117. 1871. CANTHARELLUS Adans. ex Fr., Syst. mycol. 1: 316. 1821. CHAETOCALATHUS Sing. in Lilloa 8: 518. 1942. CHAMAEOTA (W. G. Smith) Earle in Bull. New York bot. Gdn 5: 446. 1909. CHLOROPHYLLUM Massee in Kew Bull. 1898: 136. CLITOCYBE (Fr.) Kummer, Fithr. Pilzk. 26, 119. 1871. CLITOCYBULA (Sing.) Metrod in Rev. Mycol. 17: 67, 74. 1952. CLITOPILUS (Fr.) Kummer, Fuhr. Pilzk. 23, 96. 1871. COLLYBIA (Fr.) Kummer, Fuhr. Pilzk. 26, 113. 1871. CONOCYBE Fayod in Ann. Scz. nat. (Bot.) VII19: 357. 1889. COPRINUS (Pers. ex Fr.) S. F. Gray, Nat. Arrang. Brit. Pl. 1: 632. 1821. CORTINARIUS Fr., Gen. Hymen. 8. 1836. CREPIDOTUS (Fr.) Kummer, Fuhr. Pilzk. 21, 74. 1871. CRINIPELLIS Pat. inj. Bot. Paris 3: 336. 1889. CYPTOTRAMA Sing. in Lilloa 30: 375. 1960. CYSTODERMA Fayod in Ann. Scz. nat. (Bot.) VII9: 350. 1889 DELICATULA Fayod in Ann. Scz. nat. (Bot.) VII 9: 313. 1889. DESCOLEA Sing. in Lilloa 23: 256. 1950. FAYODIA Kuhner in Bull. mens. Soc. linn. Lyon 9: 68. 1930. FILOBOLETUS Henn. in Warb., Monsunza 1: 146. 1899. FLAMMULINA Karst. in Medd. Soc. Fauna Fl. fenn. 18: 62. 1891. GALERINA Earle in Bull. New York bot. Gdn 5: 423. 1909. GOMPHIDIUS Fr., Fl. scan. 339. 1835. GYMNOPILUS Karst. in Bedr. Kann. Finl. Nat. Folk 32: 400. 1879. GYRODON Opat. in Arch. Naturgesch. 2 (1): 5. 1836. GYROPORUS Quél., Ench. Fung. 161. 1886. HEBELOMA (Fr.) Kummer, Fiihr. Pilzk. 22, 80. 1871. HEBELOMINA Maire in Bull. Soc. Hist. nat. Afr. Nord. 16: 14. 1935. HEMIMYCENA (Sing.) Sing. in Rev. Mycol. 3: 194, 196. 1938. HOHENBUEHELIA Schulzer apud Schulzer, Kanitz & Knapp in Verh. zool-bot. Ges. Wien 16 (Abhand.) 45. 1866. HYGROCYBE (Fr.) Kummer, Fuhr. Pilzk. 26, 111. 1871. HYGROPHOROPSIS (J. Schroet. apud Cohn) Maire apud E. Martin-Sans, Empozs. Champ. 99. 1921. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 272 A KEY TO THE AGARICALES HYGROPHORUS Fr., Fl. scan. 339. 1835. INOCYBE (Fr.) Fr., Monogr. Hym. Suec. 2: 346. 1863. KUEHNEROMYCES Sing. & Smith in Mycologza 38: 504. 1946. LACCARIA Berk. & Br. in Ann. Mag. nat. Hist. V 12: 370. 1883. LACTARIUS DC. ex S. F. Gray, Nat. Arrang. Brit. Pl. 1: 623. 1821. LACTOCOLLYBIA Sing. in Schwezz. Z. Pilzk. 17: 71. 1939. LECCINUM S.F. Gray, Nat. Arrang. Brit. Pl. 1: 646. 1821. LENTINELLUS Karst. in Bzdr. Kann. Finl. Nat. Folk 32: 246. 1879. LENTINUS Fr., Syst. Orb. veg. 77. 1825. LEPIOTA (Pers. ex Fr.) S. F. Gray, Nat Arrang. Brit. PI. 1: 601. 1821. LEPISTA (Fr.) W. G. Smith in J. Bot., Lond. 8: 248. 1870. LEUCOAGARICUS (Locquin) Sing. in Sydowza 2: 35. 1948. LEUCOCOPRINUS Pat. J. Bot. Paris 2: 16. 1888. LEUCOPAXILLUS Boursier in Bull. Soc. mycol. France 41: 393. 1925. LIMACELLA Earle in Bull. New York bot. Gdn 5: 447. 1909. LYOPHYLLUM Karst. in Acta Soc. Fauna Fl. Fenn. 2 (1): 3. 1881. MACROCYSTIDIA Heim ex Josserand in Bull. Soc. mycol. France 49: 376. 1934. MACROLEPIOTA Sing. in Pap. Michigan Acad. Scz. 32: 141. 1948. MARASMIELLUS Murr. in N. Amer. Fl. 9: 243.1915. MARASMIUS Fr., Fl. scan. 339. 1835. MELANOLEUCA Pat., Cat. razs. Pl. cell. Tuniste 22. 1897. MELANOTUS Pat., Essat taxon. Hym. 175. 1900. MICROMPHALE Nees ex S. F. Gray em. Sing., Nat. Arrang. Brit. Pl. 1: 621. 1821. MYCENA (Pers. ex Fr.) S. F. Gray, Nat. Arrang. Brit. Pl. 1: 619. 1821. NAEMATOLOMA Karst. in Bzdr. Kann. Finl. Nat. Folk 32: 495. 1879. NOTHOPANUS Sing. in Mycologia 36: 364. 1944. OMPHALINA Quél., Ench. Fung. 42. 1886. OMPHALOTUS Fayod in Ann. Scz. nat. (Bot.) VI9: 338. 1889. OUDEMANSIELLA Speg. in An. Soc. cient. argent. 12: 24. 1881. PANAEOLINA Maire in Treb. Mus. Cz. nat. Barcelona 15 (Ser. bot. No. 2) : 109. 1933. PANAEOLUS (Fr.) Quel. in Mem. Soc. Emul. Montbeliard 11 5: 151. 1872. PANELLUS Karst. in Bzdr. Kann. Finl. Nat. Folk 32: 96. 1879. PANUS Fr., Epzcr. 396. 1838. PAXGIELOS Ei Hl SCOM TD 39 ES oO} PHAEOMARASMIUS Scherffel in Hedwigza 36: 289. 1897. PHOLIOTA (Fr.) Kummer, Fiihr. Pilzk. 22, 83. 1871. PHOLIOTINA Fayod in Ann. Sct. nat. (Bot.) VII19: 359. 1889. PHYLLOPORUS Quel., Fl. mycol. France 409. 1888. PLEUROTUS (Fr.) Kummer, Fuhr. Pilzk. 24, 104. 1871. PEUREUS) igs plaSCOTUROS 87 NS 5.oF PORPHYRELLUS Gilbert, Bolets 75, 99. 1931. PSATHYRELLA (Fr.) Quél. in Mem. Soc. Emul. Montbeliard 11 5: 152. 1872. PSEUDOBAEOSPORA Sing. in Lloydia 5: 129. 1942. PSEUDOHIATULA (Sing.) Sing. in Notula syst. Sect. crypt. Inst. bot. Acad. USSR 4 (10-12): 8: 1938. PSILOCYBE (Fr.) Kummer, Fuhr. Prlzk. 21, 71. 1871. PULVEROBOLETUS Murr. in Mycologza 1: 9. 1909. RESUPINATUS Nees ex S. F. Gray, Nat. Arrang. Brit. Pl. 1: 617. 1821. RHODOCYBE Maire in Bull. Soc. mycol. France 40: 298. 1926. RHODOPHYLLUS Quél., Ench. Fung. 57. 1886. ROZITES Karst. in Bedr. Kann. Finl. Nat. Folk 32: 290. 1879. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 A. E. WOOD 273 RUSSULA (Pers. ex Fr.) S. F. Gray, Nat. Arrang. Brit. Pl. 1: 618. 1821. SCHIZOPHYLLUM Fr., Syst. mycol. 1: 330. 1821. SIMOCYBE Karst. in Bedr. Kann. Finl. Nat. Folk 32: 416. 1879. STROBILOMYCES Berk. in Hook. J. Bot. 113: 78. 1851. STROPHARIA (Fr.) Quél. in Mem. Soc. Emul. Montbeliard 11 5: 141. 1872. SUILLUS Micheli ex S. F. Gray, Nat. Arrang. Brit. Pl. 1: 646. 1821. TRICHOLOMA (Fr.) Kummer, Fuhr. Prlzk. 25, 129. 1871. TRICHOLOMOPSIS Sing. in Schweiz. Z. Pilzk. 17: 56. 1939. TUBARIA (W.G. Smith) Gillet, Champ. France, Hym. 537. 1876. TYLOPILUS Karst. in Rev. mycol. 3 (9) : 16. 1881. VOLVARIELLA Speg. in Anal. Mus. nac. Buenos Aires 6: 119. 1899. XEROCOMUS Quél. in Moug. & Ferry, Champ. in Louis, Dep. Vosges, Fl. Vosges 477. 1887. XEROMPHALINA Kuhn. & Maire in Konr. & Maubl., Ic. Sel. JOR. (D3 B05 WEES (0)s 283. 1935. ACKNOWLEDGEMENTS Grateful acknowledgement is made to Mr. J. T. Waterhouse for helpful discussion on the manuscript. References DENNIS, R. W. G., OrTON, P. D. and Hora, F. B., 1960. — New Check List of British Agarics and Boleti — Trans. Brit. Mycol. Soc., 43, Supplement: 1-225. SINGER, R., 1975. — The Agaricales in Modern Taxonomy. 3rd Edit. Vaduz: Cramer. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 Ae = cree aN % y ns A New Species of Dakinomyiza from Queensland (Diptera: Asilidae) G. DANIELS DaniELs, G. A new species of Dakinomyza from Queensland, (Diptera: Asilidae). Proc. Linn. Soc. N.S.W. 103 (4), (1978) 1979: 275-281. Dakinomyia secuta sp. nov. is described from central Queensland, the genus previously having been recorded only from Western Australia. Prey is recorded for the new species and additional distribution records of D. froggattz (Dakin and Fordham) are noted. G. Daniels, Associate, Entomology Department, The Australian Museum, P.O. Box A285, Sydney South, Australia 2000; eee received 20 July 1978, accepted 20 Septem ber 1978. INTRODUCTION Hull (1962) records three Australian asilid genera, Opseostlengis White, Questopogon Dakin and Fordham and Dakinomyza Hardy, as being confined to the higher rainfall area of south-west Western Australia, each genus being considered as monotypic. Daniels (1976) recorded four species of Questopogon from all states except Tasmania and the Northern Territory. The known distribution of Dakznomyza is extended to Queensland where it is represented by D. secuta. Presumably collecting in inland Queensland, South Australia and Northern Territory will confirm the presence of the genus in intermediate localities. Material was examined and is housed in the following collections, abbreviated as: AM Australian Museum, Sydney ANIC Australian National Insect Collection, Canberra BM British Museum (Natural History) , London GD Author’s collection, Sydney MM Macleay Museum, Sydney WAD Western Australian Department of Agriculture, Perth. DAKINOMYIA Hardy Dakinomyza Hardy, 1934: 25. Type-species by original designation: Neosaropogon froggatta Dakin and Fordham, 1922. Flies of this genus are readily distinguished from all other Australian asilids by the distinct wing venation, the long, distal extension of the second, third and fourth posterior cells being very characteristic. They are large flies of bare aspect with a long subcylindrical and tapered abdomen. The male terminalia are - rotated 90°. The facial bristles are restricted to the dorsal epistomal margin. Hardy (1934: 25) recorded Dakinomyza secuta sp. n. from Eidsvold, Queensland under the incorrect name Neosaropogon claripennis Ricardo. An examination by Mr R. Leeke of the ‘type’ of N. claripennis at the British Museum shows that N. claripennis is not congeneric. At present N. claripennis is retained in Neosaropogon. Dakinomyza secuta sp. n. Figs 1-7 Dakinomyta claripennis (Ricardo) ; Hardy (1934: 25), misidentification. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 276 A NEW SPECIES OF DAKINOMYIA Holotype. 3, QUEENSLAND: Blackdown Tableland, Expedition Range, 6.i.1976, G. Daniels, (AM). Paratypes. QUEENSLAND: 14 6, 10 , same data as holotype except 6- 9.1.1976 (1d, 12, BM; G.D); 1 9, same data as holotype except 23.xii.1972, M. S. Moulds, (G.D.) ; Eidsvold, 11.iv.1924, x.1929-iv.1930, ii-vi.1923 (46,29, AM,24d, 22, ANIC), C. Gibbons, T. L. Bancroft (1 2 from Eidsvold identified as Neosaropon claripennis in G. H. Hardy’s handwriting) . Other Material Examined (Non-Type) : Eidsvold, no data, (1d, 19 AM,14d,12 ANIC); 1d, Eidsvold, ii-vi.1923, Mackerras (MM). MALE. Head. Frons, face, occiput and vertex pale yellow pollinose. Mystax comprised of 2 or 3 rows of erect, stout, pale yellow bristles confined to dorsal margin of epistoma. Fronto-orbital bristles absent but a group of 4-8 stout, admixed black and yellow bristles present on lower anterior corner of ocellar tubercle. Ocellar bristles confined to apex of occéllarium, comprised of 3 or 4 pair of long stout black bristles with several weaker scattered elements. Palpi brownish, white haired; second distal segment with laterally offset apical pore. Proboscis black, with a laterally compressed dorsal ridge. Occipital bristles pale yellow, more or less confined to a single row with some weaker elements along each side. Antennae orange; first and second segments dusted yellow, setae black; third segment with an apical pit at an oblique angle on inner margin, the pit bearing a short blunt concealed spine. Relative lengths of antennal segments 1:1.05-1.15:3.14-3.86. Thorax. Mesoscutum brownish with broad blackish medial line and pre- and postsutural dorsocentral areas, the latter sometimes narrowly joined dorsally. Scutellum brownish. Pleura grey dusted. Pronotum with a row of moderately stout yellowish bristles with a posterior and anterior row of weak erect bristles. Posterior pronotum laterally with 2-7 strong erect long black bristles surrounded by weaker white elements. Mesoscutum with abundant short appressed black setae. The medial setae are in a double row and have a bare submedial line either side. Dorsocentral areas bare except for a few anterior setae on dark presutural mark. Dorsocentral bristles not extending anteriorly to suture, posterior elements incurved. Humeral callus with 5 or 6 bristles anteriorly, mixed black and yellowish; posterior surface setate. 5-7 strong black presutural, 3 supra alar and 2 or 3 postalar bristles. Scutellar disc bare; margin with a pair of long, black convergent bristles. Metanotum strongly bulbous; micropubescent only. Mesopleuron, pteropleuron and anepisternite micropubescent. Pleurotergite with a double row of fine bristles, dorsally being finer and denser, sometimes forming a ‘tuft’. Metapleuron with some weak bristles posteriorly. Legs. Orange-brown. Fore and mid femora with dense appressed black setae and only a single short subapical bristle posteriorly on fore femur and 1 or 2 similar bristles on mid femur. Fore tibia with a row of 2-4 antero- and 5 or 6 posterodorsal short bristles; a row of 6 posteroventral bristles, short except for a single long stout bristle at apical third, usually being the fifth bristle in the row; apically with a fan of 6 or 7 bristles of varying lengths. Tibia dark brown on apical fourth. First 3 tarsal segments yellow, with dense appressed setae; third segment occasionally deep brown on apical third; segments 4 and 5 deep brown; basitarsus twice as long as succeeding segment. Mid tibiae with dense short appressed setae; dark brown on apical fourth; a row of 4 to 6 short dorsal bristles; 3 evenly spaced, long, anterodorsal bristles; a row of 3-5 short anteroventral bristles; 2 long ventral bristles arising from mid point and apical fourth of tibia; a row of 4 or 5 anteroventral bristles. Mid tarsus similar to fore tarsus. Hind femur with dense appressed setae, deep brown on apical sixth; a weak bristle present anteriorly on mid point of femur. Hind tibia yellow, deep brown on apical third; with dense short appressed black setae. A row of 4 long dorsal bristles, 3 long anterodorsals, 2 being before basal half and one on apical third; a row Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 G. DANIELS 277 Figs 1-7. Dakinomyia secuta sp.n., terminalia: (1) tergites of male, ventral view; (2) sternites of male, dorsal view; (3) aedeagus, lateral view; (4) aedeagus, ventral view; (5) tergites of female, dorsal view; (6) sternites of female, dorsal view; (7) spermathecae and furca, dorsal view. Setae are omitted from Figs 1,2,5and6. c=cerci, p= proctiger, s = sternite, t = tergite. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 278 A NEW SPECIES OF DAKINOMYIA of 4 or 5 anteroventrally. Hind tarsus deep brown; first and second segments yellow on basal sixth and half respectively. Basitarsus about 2.5 times as long as succeeding segment. Wings. Venation and shape similar to D. froggattzz as illustrated in Hull, 1962 fig. 510. Abdomen. Strongly tapered; orange-brown. First tergite mostly black, narrowly orange-brown along posterior margin. Second tergite blackish brown on basal fourth, this same area being white pruinescent. White pruinescent stripe on lateral margin of tergites 1-7. Fine short appressed setae present on tergites. Lateral margin of tergites 1-4 with fine pile, relatively long and dense on first tergite and becoming shorter and less dense on each succeeding tergite. Sternites white pruinescent with fine white erect setae. Segment 8 orange, recessed into preceeding segment. Termznalza (Figs 1-4) black, rotated 90°. FEMALE. Similar to male except as follows: Abdomen. Tergites 5-8 shining, with sparse, erect white setae. Tergites 7 and 8 black. Tergites 1-5 with lateral white pruinescent line. Termznalza (Figs 5 and 6) black. Acanthophorites with 6 or 7 pairs of long, stout, blunt spines. DIMENSIONS: length of body, d 26.0 - 30.1 mm, @ 26.5 - 32.5 mm; length of thorax, 5 6.5-7.4mm; 9 6.8 - 7.7 mm; length of wing, d 17.5 - 19.2mm; 9 17.5 - 19.7 mm. MORPHOLOGY OF THE FEMALE TERMINALIA The female terminalia, as herein discussed, comprise the genital and postgenital segments (abdominal segments 8-11) . SEGMENT EIGHT Tergite 8 (Fig. 5). On the expanded female abdomen, tergite 8 (t8) is a medium sized, easily observable sclerite and is relatively unmodified. The anterior margin is convex, a similar condition existing on the posterior margin, although less strongly so. The posterior margin is somewhat membranous centrally. The ratio of medial length to maximum width is 3.7 to 6.8. Sternite 8 (Fig. 6). Like t8, sternite 8 (s8) is a relatively large and conspicuous sclerite although it is a more complex structure than t8 and has undergone diverse modifications. A medial recess is present and the submedial area is extended posteriorly. The posterior margin has a submedial lobe extending along the inner surface of s8. SEGMENT NINE Tergite nine (and tergite ten) (Fig. 5). Tergite 9 (t9) and tergite 10 (t10) cannot be entirely separated as they appear to be fused. t9 would appear to be reduced to asmall narrow, double crescent shaped sclerite at the anterior margin of the deeply, medially divided t10, with a narrow pointed medial extension posteriorly. Sternite nine (Fig. 6). Sternite 9 (s9) appears as a small sclerite, apparently fused to each of the lateral, anteriorly directed, posterior margins and extends anteriorly and medially to a point slightly posterior to the posterior margin of the furca. Each sclerite is concave posteriorly and more or less sharply angulate anteriorly. SEGMENT TEN Tergite ten (Fig. 5). t10 is a rounded sclerite medially divided posteriorly, the division extending towards the anterior margin but not reaching it. Each acanthophorite, as this divided tergite is generally referred to, possesses strongly modified setae, usually six in number but occasionally seven. | Sternite ten (? plus eleven) (Fig. 6). Sternite 10 (? plus 11) (s10) is a broad and narrow membranous sclerite that partially overlaps the medial extension of s8. Each anterior lateral corner overlies part of s9. The medial lateral margin appears to be joined with the ventral margin of s10. Proc. LINN. Soc. N.S.W., 103 (4), (1978) 1979 G. DANIELS 279 CERCI (Fig. 5). The cerci are attached to the posterior margin of t10 and bear a row of blunt spine-like bristles along the extreme posterior margin. The remaining ventral surface is densely covered with short, fine, hair-like bristles. FEMALE REPRODUCTIVE SYSTEM The female reproductive system comprises a pair of tubular accessory glands on long slightly narrower ducts (Fig. 7), 3 elongate spermathecae with brownish vesicles, the spermathecal ducts being approximately twice the diameter of the spermathecae and have a short common duct before entering the median oviduct. Viewed dorsally the furca is semicircular with a vertical rib on the dorsal surface. This rib is concave laterally and is flared anteriorly. The ovaries (omitted from Fig. 7) are long and narrow, extending from the posterior margin of segment 1 to the anterior margin of segment 8 and have a short oviduct. MORPHOLOGY OF THE MALE TERMINALIA The male terminalia comprise the eighth to eleventh abdominal segments. SEGMENT EIGHT Tergite eight (Fig. 1). t8 is a medium sized, easily observable sclerite and has undergone some modification. The anterior margin is concave whilst the posterior margin is convex and tends to form a lobe medially. The ratio of medial length to maximum width is 4.8 to 9.3 Sternite eight (Fig. 2). s8 is a small relatively inconspicuous sclerite, and is rectangular in shape with the posterior margin slightly convex. SEGMENT NINE Tergite nine (epandrium) (Fig. 1). t9 is slightly longer than t8 and fully cleft medially, the two halves rounded, obtuse and converging posteriorly. The dorsal margin of each half is rounded, forming a small lobe. The anterior margin is straight but angled to the medial axis. Sternite nine (hypandrium) (Fig. 2). s9 is a well-developed sclerite with a distinctly convex anterior margin. The lateral margins are narrowed posteriorly and the posterior margin has a distinct medial projection. SEGMENT TEN Tergite ten (Fig. 1). t10 is reduced to a pair of sclerites lying below the posterior margin of t9. The anterior margin is at approximately 45° to the medial line and slightly concave. The anterolateral margin is extended into a narrow lobe. The lateral margins are almost straight with a convex lateral projection towards anterior margin. Medially, each sclerite has a distinct recess just behind the convex posterior margin. Sternite ten (coxite) (Fig. 2). sl0 appears as a pair of short curved processes (‘styles’) which arise subapically from the inner surface. The ventral style has parallel sides for most of its length, being bluntly rounded apically and slightly enlarged basally. The dorsal style is much stouter than the ventral style, generally about 3 times as wide. At about one fourth of the length of this process a small dorsal spur is present. PROCTIGER The proctiger (Fig. 1) is a rectangular membranous sclerite with the anterior margin slightly wider than the posterior margin, extending slightly beyond s10 and has an indistinct medial crease. The posterior margin is almost straight with a small setose lobe on each lateral corner and a broad indistinct convex lobe medially. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 280 A NEW SPECIES OF DAKINOMYIA AEDEAGUS The aedeagus (Figs 3 and 4) is short, widened anteriorly and sulcate posteroventrally. The anterior dorsal margin and a lateral ventral groove are membranous. D. secuta differs from D. froggattzz in the following characters: D. froggatti has 3rd antennal segment brownish; thorax with dorsum uniformly brown-black; lst abdominal tergite black dorsally on basal third and on lateral margin sublaterally tergites 2-5 with a black line, extreme margin orange-brown, with a yellow pruinescent line, tergites 6-8 black orange-brown lateral margin; sternites deep black-brown; femora red-brown, dorsally black, and apically widened tibia with, at most, basal third yellow; coxae orange-brown to deep brown; first tergite with black microtrichiae dorsally, white haired laterally, and with 4-5 black stout bristles laterally; tergites two to five with extremely fine, stout orange microtrichiae. FORAGING AND PREDATION Observations at Expedition Range, Queensland While awaiting prey, the asilids rest on rocks, soil and sticks close to the ground, though the greatest preference is for large, bare rocks. The flies land on sunny areas and if even partly covered by shadow will usually move immediately. Upon alighting, these flies take up a characteristic position with their bodies turned perpendicularly to the sun. The bodies were normally held high off the substrate with the tip of the abdomen held slightly above the substrate. Predation was observed mostly in the morning from about 7.00 am to 10.30 am. When prey is sighted, the predator’s whole body turns to face it. Forage flights were usually short, ranging from about 30 cm to a metre. Prey was taken only in the air and was usually impaled upon being captured. After capturing prey, the asilid would fly up to 3 m to find a suitable perch to manipulate prey, either a grass stem or twig about 30-45 cm above the ground. Upon alighting one of the fore legs was used as an anchor whilst the other fore leg and the middle legs manipulated the prey. However if a fly was disturbed whilst feeding it would seek a large bare area of rock or soil and continue feeding. Prey selection is presented only to generic level as specific identifications are not available. It is noteworthy that the prey of all female asilids captured were Hymenoptera and three of the Hymenoptera were males. Prey taken: 1 9 with Ropalidea sp. (Hymenoptera: Vespidae) 1 d with Rhytzdoponera sp. (Hymenoptera: Formicidae) 1 d with nr Dasypogon sp. (Diptera: Asilidae) 1 2 with Apis mellifera (L.) (Hymenoptera: Apoidea) 1 2 with Tachytes sp. (Hymenoptera: Sphecidae) 1 2 with Campsomeris sp. (Hymenoptera: Scoliidae) 1 d with Hemiptera: Cicadidae 1 d with Diptera: Asilidae (undetermined genus) Dakinomyva froggattz (Dakin and Fordham) Neosaropogon froggatti# Dakin and Fordham, 1922: 523. Dakinomyta froggatta (Dakin and Fordham) — Hardy 1934: 25. The only recorded locality for this species is the type locality, Bremer Bay, Western Australia. The following specimens extend the distribution approximately Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 G. DANIELS 281 430 km to the north east: 1 9 Lake Cronin, W.A. 16.iii.1970 K. T. Richards (WAD) 1 9 Bakers Hill W.A. 24.ii.1969 T. Burbidge (WAD) ACKNOWLEDGEMENTS Thanks are due to the Queensland Forestry Department for permission to traverse and collect in land under its jurisdiction; to Mr J. Hodges, Macleay Museum for allowing access to material; Mr M. S. Moulds for the presentation of material; to Mr K. T. Richards and Dr D. H. Colless for loan of material; Mr G. A. Holloway for determination of Hymenoptera and Mr R. Leeke for examination of type material in the British Museum. References Dakin, W. J. and ForpHa, M. G. C., 1922. — Some new Asilidae from Western Australia. Ann. Mag. nat. Hist., (9) 10: 517-530. DaniELs, G., 1976. — Three new species of Questopogen Dakin and Fordham (Diptera: Asilidae) from Australia. Proc. Linn. Soc. N.S.W., 100: 223-230. Harpy, G. H. H., 1934. — The Asilidae of Australia. — Part I. Ann. Mag. nat. Hist., (10) 13: 498-525. Hutt, F. M., 1962. — Robber flies of the world: the genera of the family Asilidae. U.S. Nat. Mus. Bull. 224. Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 Acari Acarina: Phytoseiidae Agaricales, a key to the Australian genera Alcyonaria from a temperate shallow water reef Algae from a temperate shallow water reef Amblysetus lazlae, sp. nov. Amblyseius neolentiginosus, sp. nov. Amblyseius neovictorzensts, sp. nov. Amphipoda from a temperate shallow water reef Anderson, D. T., The Sir William Macleay Memorial Lecture 1978. Natural History today Annelida: Oligochaeta Anura: Hylidae Ash, S. R., see Holmes, W. B. K. Australofenestella brookeri, gen. & sp. nov. Australofenestella cincta, gen. nov. Australofenestella(?) keepitensis, sp. nov. Australofenestella macleayenszs Australofenestella malchi Australofenestella trevallynensis, gen. & sp. nov. Australofenestella stroudensts minuta, gen. & subsp. nov. Australofenestella stroudenszs Australofenestella stroudensis stroudensts Australopolypora altinodosa, gen. nov. Australopolypora keppelenszs, gen. nov. Australopolypora keppelenszs parvula gen. & subsp. nov. Australopolypora neerkolenszs Australopolypora palenensis Australopolypora rawdonensis, gen. & sp. nov. Australopolypora scalpta Bass, A. J., Records of little-known sharks from Australian waters Bell, G. D., see Webb, J. A. Biota of a temperate shallow water reef, by J. E. Watson Brachiopods, late Ordovician articulate, from Gunningbland Bryozoa from a temperate shallow water reef Bryozoans, fenestrate, with large apertural form in the Carboniferous of eastern Australia Carboniferous fenestrate bryozoans Centroscymnus crepidater, first record from Australian waters Centroscymnus owstonz, first record from Index VOLUME 103 Page 43, 189 Australian waters 217 ~=Chlamydoselachus anguineus, first record from Australian waters 275 Cirrhigaleus barbifer, first record from Australian waters 229, 234 Cladophlebis carne, sp. nov. Cladophlebis cf. mendozaensts, Triassic, 228, 233 from the Lorne Basin 217 ~ Cladophlebis sp. indet., from the Lorne 222 ~—- Basin 222 Clathrodictyid stromatoporoids, Ordovician 229, 235 Clathropteris sp. Cliefdenellid stromatoporoids, Ordovician ?Coniopteris burejenszs, from the Lorne 1 Basin 37 Coniopteris? ramosa, sp. nov. 23 Crustacea, Caridea, Atyidae Crustacea: Conchostraca Culictnomyces, an insect pathogenic 154 fungus in the Sydney area Mee Dakinomyia froggatti, notes on its 164 distribution 16] Dakinomyra secuta, sp. nov. 160 Daniels, G., A new species of Dakinomya from Queensland (Diptera: Asilidae) 158 Debenham, M. L., see Russell, R. C. Dermanyssid mites (Acari), mostly from 157 Australasian rodents, by R. Domrow 155 Dicroidium dubium var. australe, from 155 the Lorne Basin 146 Dicroidium voiseyt, Sp. Nov. 149 Dicroidium zuberi var. feistmantelliz, from the Lorne Basin 150 Dictyophyllum acutilobum 1a Dictyophyllum sp. cf. D. acutilobum les Dictyophyllum bremerense Dictyophyllum davidii Ise Diplotrema tyagarah, sp. nov. 149 Dipteridaceae, review of Australian, by R. Herbst 947 Dolerozdes sp., Ordovician, from Gunningbland Domrow, R., Some dermanyssid mites 227 (Acari), mostly from Australasian rodents 175 Domrow, R., see Fain, A. Dyne, G. R., A new species of 229, 234 Microscolex (Diplotrema) (Annelida: Oligochaeta) from New South Wales Echinonyssus butantanenszs, new data 135 Engel, B. A., Fenestrate bryozoans with 135 large apertural form in the Carboniferous of eastern Australia 252 Equisetaceous stems from the Lorne Basin Page 252 247 250 189 37 204 135 51 Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 284 Eulaelaps stabularis, new data Eunaptus sinensis Fain, A., & Domrow, R. The family Hypoderidae (Acari) in Australia Feeding habits and structure of the gut of the Australian freshwater prawn Paratya australiensts Kemp (Crustacea, Caridea, Atyidae), by P. Gemmell Fenestrate bryozoans with large apertural form in the Carboniferous of eastern Aus- tralia, by B. A. Engel Final larval instar of Phaenocarpa (Asobara) persimilis Papp (Hymenoptera, Braconidae, Alysiinae) from Australia, by J. R. T. Short Freshwater sponges from the Northern Territory (Porifera: Spongillidae), by J. Stanisic Gartside, D. F., see Martin, A. A. Gemmell, P., Feeding habits and structure of the gut of the Australian freshwater prawn Paratya australzensis Kemp (Crustacea, Caridae, Atyidae) Granite Belt in southern Queensland and northern New South Wales, a new species of Limnadia from Gunningbland, N.S.W., late Ordovician articulate brachiopods from Gut, structure of the, in Paratya australienss Haemolaelaps domrowi Haemolaelaps penelope Halarachne mzroungae Hausmannia (Hausmannia) bulbaformis Hausmannia (Hausmannia?) wilkinsi Hausmannia (Protorhipis) sp. cf. H. (P.) deferrarzisc Hemipristis elongatus, first record from Australian waters Herbst, R., Review of the Australian Dipteridaceae Hexanchus griseus Holmes, W. B. K., & Ash, S. R., An early Triassic megafossil flora from the Lorne Basin, New South Wales Holothuroidea from a temperate shallow water reef 230, Hydroids from a temperate shallow water reef 228, Hymenoptera, Braconidae, Alysiinae Infurca tessellata, gen. & sp. nov. Isopoda from a temperate shallow water reef Jensen, Hans Laurits (1898-1977). Memorial Series No. 26 Karibacarpon feistmantellz, sp. nov. Key to the Australian genera of the Agaricales, by A. E. Woods Labechiid stromatoporoids, Ordovician Laelaps aella Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 INDEX 204. ~=Laelaps assimilis 195 128 Laelaps bycalia, sp. nov. 199 Laelaps cybiala 203 43 Laelaps echidninus 196 Laelaps janalis, sp. nov. 197 Laelaps lybacia, sp. nov. 203 Laelaps nuttalli 195 999 ~=~—Laelaps pammorphus 197 Laelaps rothschildz 196 Laelaps southcotti 194 135 Laelaps sp. 194 Laelaps wasselli 195 Late Ordovician articulate brachiopods from Gunningbland, central western New 17] South Wales, by I. G. Percival 175 Lee, D. J., see Russell, R. C. Lepidopterts madagascariensis, from the 123 Lorne Basin 64 aff. Leptellzna sp., from Gunningbland 181 Limnadza urukhat, sp. nov. 239 Linnean Society of New South Wales. A key to the Memorial Series (1928-1978) 133 Linnean Society of New South Wales. 209 Record of the Annual General Meeting 1978. Reports and Balance Sheets 75 Litorza peronzz Complex (Anura: 237 Hylidae), a new species from eastern Australia 23 175 Litoria tyler?, sp. nov. 25 Littlejohn, M. J., see Martin, A. A. 209 Loftus-Hills, J. J., see Martin, A. A. 191 Lorne Basin, N.S.W., Triassic megafossil 192 = flora from 47 207. _—_ Lycopod stems from the Lorne Basin 51 19 Macleay Memorial Lecture 1978, by 18D. T. Anderson: Natural History today 1 Martin, A. A., Watson, G. F., Gartside, 18D. F,, Littlejohn, M. J., & Loftus-Hills, J. J., A new species of the Lztoria peronz 248 Complex (Anura: Hylidae) from eastern Australia 23 7 Memorial Series No. 26. Hans Laurits 248 Jensen 1898-1977 131 Memorial Series of the Linnean Society of New South Wales (1928-1978), a key 133 47 Mesolaelaps australiensis 189 Mesolaelaps bandicoota 190 235 Metania ovogemmata, sp. nov. 124 Microphyllopteris sp., from the Lorne ze Basin 58 171 Microscolex (Diplotrema) (Annelida: 186 Oligochaeta), a new species from N.S.W. 37 Microscolex ‘(Diplotrema) tyagarah, sp. 230 ~—s nov. 38 Mites, dermanyssid 189 131 Mites, hypoderid 43 Mollusca from a temperate shallow water 60 reef 229, 234 obs Natural habitat of the insect pathogenic fungus Culicznomyces in the Sydney area, 85 by R. C. Russell, M. C. Debenham & 196 D. J. Lee 71 INDEX 285 Natural History today. Macleay Memorial Lecture 1978, by D. T. Anderson 1 Neottzalges (Pelecanectes) evanst 43 Neottialges (Pelecanectes) plegadicola 54 Neottialges (Pelecanectes) tasmaniensis, sp. nov. 44 New species of Dakznomyza from Queensland (Diptera, Asilidae), by G. Daniels 275 New species of Lemnadza (Crustacea: Conchostraca) from the Granite Belt in south Queensland and northern New South Wales, by J. A. Webb & G. D. Bell 237 New species of Lztorta peronit Complex (Anura: Hylidae) from eastern Australia, by A. A. Martin, G. F. Watson, D. F. Gartside, M. J. Littlejohn & J. J. Loftus-Hills 23 New species of Microscolex (Diplotrema) (Annelida: Oligochaeta) from New South Wales, by G. R. Dyne 37 Nodal diaphragms (Sphenopsida) from the Lorne Basin 53 Northern Territory, freshwater sponges from 123 Oepikina? sp., from Gunningbland 183 Ophiuroidea from a temperate shallow water reef 230, 235 Ordovician articulate brachiopods from Gunningbland 175 Ordovician stromatoporoids. Presidential Address by B. D. Webby 83 Ornithonyssus bacoti 207 Ornithonyssus syluarum 207 Paratya australiensis, feeding habits and structure of the gut 209 Peramelaelaps bandicoota 192 Percival, I. G., Late Ordovician articulate brachiopods from Gunningbland, central western New South Wales 175 Phaenocarpa (Asobara) persimilis, the final larval instar of 171 Polychaetes from a temperate shallow water reef 229, 234 Porifera: Spongillidae 123 Presidential Address 1978, by B. D. Webby. The Ordovician stromatoporoids 83 Pteruchus barrealensis var. feistmantellzt, comb. et var. nov. 62 Radiospongilla streptasteriformis, sp. nov. 126 Reef, biota of a temperate shallow water, (at 34° 01’ 20" S, 151° 14’ 20" E) 227 Review of the Australian Dipteridaceae, by R. Herbst 7 Rhipidopsis? narrabeenenszs, from the Lorne Basin 65 Russell, R. C., Debenham, M. L., & Lee, D. J., A natural habitat of the insect pathogenic fungus Culicenomyces in the Sydney area 71 Scaphorthis? aulacis, sp. nov. 177 Schicha, E., Three new species of Amblysezus Berlese (Acarina: Phytoseiidae) from Australia 217 Sharks, little-known, from Australian waters 247 Short, J. R. T., The final larval instar of Phaenocarpa (Asobara) persimilis Papp (Hymenoptera, Braconidae, Alysiinae) from Australia 171 Skilliostrobus australis, from the Lorne Basin 50 Sowerbyella antictpata, sp. nov. 182 Sponges, freshwater, from the Northern Territory 123 Sponges from a temperate shallow water reef 228, 233 Spongillids, origins and relationships of the Australia 128 Stanisic, J., Freshwater sponges from the Northern Territory (Porifera: Spongillidae) 123 Stromatoporoids, Ordovician, origins and interrelationships 112 Stromatoporoids, The Ordovician. Presidential Address by B. D. Webby 83 Stromatoporoids, Ordovician, stratigraphic distribution 104 Taeniopteris lentriculiforme, from the Lorne Basin 64 Tchan, Y. T., & Vincent, J. M., Hans Laurits Jensen 1898-1977. Memorial Series No. 26 131 Thaumatopterzs shirley, sp. nov. 16 Three new species of Amblysezus Berlese (Acarina: Phytoseiidae) from Australia, by E. Schicha 217 Triassic megafossil flora from the Lorne Basin, N.S.W. 47 Trichosurolaelaps crassipes 205 Trichosurolaelaps striatus 206 Vallance, T. G., A key to the Memorial Series of the Linnean Society of New South Wales (1928-1978) 133 Vincent, J. M., see Tchan, Y. T. Voltziopsis townrovit, sp. nov. 65 Watson, G. F., see Martin, A. A. Watson, J. E., Biota of a temperate shallow water reef 227 Webb, J. A., & Bell, G. D., A new species of Lzmnadia (Crustacea: Conchostraca) from the Granite Belt in southern Queensland and northern New South Wales 237 Webby, B. D., Presidential Address 1978. The Ordovician stromatoporoids 83 Wood, A. E., A key to the Australian genera of the Agaricales 255 Proc. Linn. Soc. N.S.W., 103 (4), (1978) 1979 ADVICE TO AUTHORS The Linnean Society of New South Wales publishes in its Proceedings original papers and review articles dealing with biological and earth science. 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Authors are urged to supply such photographic reproductions which, if made to a scale appropriate to the size of a printed page, will show whether ornament and lettering can be read in the final print. All photographs, whether for half-tone or line illustrations, should be high-contrast, glossy prints. Each illustration should be identified (author’s name, Fig. no. and orientation) in pencil on the back. . Tables should be submitted in a clear format on separate sheets. Like illustrations, they should be designed to fit a single page of the journal. PROCEEDINGS of the LINNEAN SOCIETY OF NEW SOUTH WALES VOLUME 103 Issued 28 December 1979 i ee CONTENTS: 135 OT: 175 189 * PART 3 B. A. ENGEL Fenestrate Bryozoans with Large Apertural Form in the Carboniferous of eastern Australia J. R. T. SHORT -The Final Larval Instar of Bhaenocdrpa Coe persimilis Papp Veni ake Braconidae, Alysiinae) from Australia |. G. PERCIVAL Late Ordovician Articulate Brachiopods from eon abla Central Western New South Wales PART 4 R. DomROW Some Dermanyssid Mites (Acari), mostly from Australasian Rodents P. GEMMELL Feeding Habits and Structure of the Gut of the Australian Freshwater Prawn Paratya australiensis Kemp (Crustacea, Caridea, Atyidae) - E. SCHICHA Three New Species of Amblvcenus Berlese (Acarina: Phytoseiidae) from Australia . J. E. WATSON Biota of a Temperate Shallow Water Reef J. A. Wess and G. D. BELL A New Species of Limnadia (Crustacea: Bopcng cea) from the Granite Belt in Southern Queensland and Northern New South Wales ‘A. J. Bass Records of Little-kKnown Sharks from Australian Waters A. E. Woop A Key to the Australian Genera of the Agaricales G. DANIELS A New Species of Dakinomyia from Queensland (Diptera: Asilidae) Index to Volume 103 Printed by Southwood Press Pty Limited, 80-92 Chapel Street, Marrickville 2204 -MBL/WHOI LIBRARY _ | Il LAET H eh svi Lee ts Pinte ats Hagadbat ae ace Re tara it seit Aaa ig te wy Pree iis May tebe ngs we ab Riaad Sead actaset se) Fel dgey Sith away 5; saat ati wits etlt Tyee ee y Telateticn meh ee ae as S teat ast HA pe ets Pagee joys