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' Ae i AE uw AeA . eit A us \s ren Rear eae > vi d : F ee SP A : : any che Nts Srl geit Wig 6: BEY an Spe. arian tee ~ ae ; Ady Wath Awe a vapedivges Wein gts pega Ents Pies A Hk siabas 9 haat. eee shiv. be beh Cae : Wray ‘ ; or te t 2 he Rug coed hehe i : iby ‘ Mi “afin ail . ‘ id : Hu tae Ur ‘NOR a ; te ‘ } : \ rho } ae 2 Suey p ied ye gases ar Ai fn AB regs Fee tt ; ‘ ‘ t ‘ & hoa te " eas , Ped fa EF 3 Ata be ‘ ( i Soni 7 wr ae pete iy yep Neh tett Pe sor Hy “i Baa Sota Si inaie. den eh bese fStop ae a fe thE PGs Path Hit bh ey f it Hes u i { he afkcvalet iis 4 ay 9948: 101 Ae Proceedings of the. to) °° x Te Linnean Society of New South Wales VOLUME 97 No. 429-432 CONTENTS OF PROCEEDINGS, VOLUME 97 PART 1 (No. 429) (Issued 22nd June, 1972) CONTENTS Page Annual General Meeting : Report on the Affairs of the Society for the Year .. sa ae 1 Elections .. Se ane wi a ee a8 e: aa 5 Balance Sheets ad is hg wt ata gine fe ote 6 JOHNSON, L. A. S. Presidential Address: Evolution and Classification in Hucalyptus ae rs Ha se = ee ig ss aa DARTNALL, ALAN J. A Brooding Echinoid from Tasmania .. = dO) KHAN, A. G., and VALDER, P.G. The Occurrence of Root Nodules in the Ginkgoales, Taxales, and Coniferales. (Plate 1) ai oe a 35 PONDER, W. F., and STANBURY, P. J. Type Specimens in the Macleay Museum, University of Sydney .. ws 3 a Pe si 0 SAD SANGHI, A. K., and BAKER, E. P. Genetic Bases for Resistance in Two Common Wheat Cultivars to Stem Rust Strains of Unusual Avirulence 56 TRACEY, D. J. LEvisceration and Regeneration in Thyone okeni (Bell, 1884) ats oe ae mee Ne a os 5 one (74 TYLER, MIcHAEL J., MARTIN, ANGUS A., and WATSON, GRAEME F. A New Species of Hylid Frog from New South Wales. (Platem) .. 82 WALKER, J.C. A Fellodistomid Cercaria from Mytilus planulatus So Old Notes and instructions for authors .. a: ae a oe we 93 PART 2 (No. 430) (Issued 11th October, 1972) CONTENTS BASDEN, RALPH. A Series of Oligosaccharides, Occurring in the Honeydew of Insects, Based on Turanose .. Browne, W. BR. Grey Billy and Its Associates in Eastern Australia RICHARDSON, LAURENCE R. A New Genus Based on the Seven-banded Richardsonianus dawbini Richardson 1969 (Hirudinoidea: Richard- sonianidae) .. SELKIRK, D. R. Fossil Manginula-like Fungi and Their Classification. (Plates M1-vm) WEBBY, B. D. The Rugose Coral Palaeophyllum Billings from the Ordovician of Central New South Wales. (Plates v1, Ix) Notes and instructions for authors Page 95 98 130 141 150 159 PART 3 (No. 431) (Issued 8th January, 1973) CONTEN'S FAIn, ALEX, and DomMrRow, ROBERT. Two New Fur-Mites (Acari: Atopomelidae) from an Australian Tiger Cat (Marsupialia: Dasyuridae) Hamonp, R. Four New Copepods (Crustacea: Harpacticoida, Canuel- lidae) Simultaneously Occurring with Diogenes senex (Crustacea: Paguridea) near Sydney .. Price, InFRYN. A New Permian and Upper Carboniferous (?) Succession near Woodsreef, N.S.W., and Its Bearing on the Palaeogeography of Western New England SippiqI, M. Y., CAROLIN, R. C., and ANDERSON, D. J. Studies in the Ecology of Coastal Heath in New South Wales. I. Vegetation Structure SLATYER, R. O. Sir William Macleay Memorial Lecture, 1972. Energy Flows in the Biosphere—the Impact of Man Notes and instructions for authors Page 161 165 211 225 237 PART 4 (No. 432) ba rai (Issued 30th April, 1973) CONTENTS Page Kort, Patricia. Notes on Some Ascidians from Port Jackson, Botany Bay and Port Hacking, New South Wales 241 Kortt, Patricia. Plurellidae, a New Phlebobranchiate Family of the Ascidiacea 258 KRATOCHVIL, M., HANNON, NOLA J., and CLARKE, LESLEY D. Mangrove Swamp and Salt Marsh Communities in Southern Australia. (Plate x) 262 Morrat, LYNETTE A. The Concept of Primitiveness and Its Bearing on the Phylogenetic Classification of the Gekkota 275 RENBUSS, Marya A., CHILVERS, G. A., and Pryor, L. D. Microbiology of an Ashbed. (Plate x1) 302 Abstract of Proceedings oll List of Members 316 List of Plates 324 List of New Species 324 Index 325 Notes and instructions for authors 333 ane is 6 to J ale: ols es : i Ms si ine Ae Hi Proceedings of the ‘Linnean Society of New South Wales Issued 22nd June, 1972 VOLUME 97 PART | No. 429 The binscan Society of New South Wales Founded 1874. Incorporated 18384 ‘“‘ For the cultivation and study of the science of Natural History in all its branches ”’ OFFICERS AND COUNCIL, 1972-73 President H. G. Cogger, M.Sc., Ph.D. Vice-Presidents L. A. 8S. Johnson, D.Se.; T. G. Vallance, B.Sc., Ph.D. ; N. G. Stephenson, M.Sc., Ph.D. ; R. C. Carolin, B.Sc., Ph.D., A.R.C.S. Honorary Treasurer Joyce W. Vickery, M.B.E., D.Sc., F.L.S. Secretary Mrs. Ruth J. Inall Counce D. J. Anderson, Ph.D. Elizabeth C. Pope, M.Sc., C.M.Z.S. D. T. Anderson, D.Sc. P. J. Stanbury, Ph.D. R. K. Bamber, F.S.T.C. N. G. Stephenson, M.Sc., Ph.D. Barbara G. Briggs, Ph.D. F. H. Talbot, M.Sc., Ph.D., F.L.S. W. R. Browne, D.Sc., F.A.A. E. LeG. Troughton, F.R.ZS8. R. C. Carolin, B.Sc., Ph.D., A.R.C.S. P. G. Valder, B.Sc.Agr., Ph.D. H. G. Cogger, M.Se., Ph.D. T. G. Vallance, B.Se., Ph.D. Mary M. Hindmarsh, B.Sc., Ph.D. Joyce W. Vickery, M.B.E., D.Sc., F.L.S. L. A. S. Johnson, D.Se. G. P. Whitley, F.R.Z.S. Auditors W. Sinclair and Company, Chartered Accountants Linnean Macleay Lecturer in Microbiology, University of Sydney K. Y. Cho, Ph.D. Linnean Macleay Fellow of the Society as from Ist July, 1971 Mrs. Lynette Anne Moffat, B.Sc.Hons. The Society’s Headquarters are in Science House, 157 Gloucester Street, Sydney, N.S.W. 2000, Australia ANNUAL GENERAL MEETING 29th MARcH 1972 The Ninety-seventh Annual General Meeting was held in the Society’s Rooms, Science House, 157 Gloucester Street, Sydney, on Wednesday, 29th March 1972, at 7.30 p.m. Dr. L. A. 8. Johnson, President, occupied the chair. The minutes of the Ninety-sixth Annual General Meeting (31st March 1971) were read and confirmed. REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 1971 Publication The Society’s Proceedings for 1970, Part 3, was published on the 31st March 1971, and for 1971, Parts 1, 2 and 3, on the 17th June, 15th September and 10th December 1971. Donations totalling $250.00 were made during the year towards the cost of publication, which was as follows: 5) ) Block-making 3 874.92 Donation... ve 250.00 Printing y .. 93,629.60 Subscriptions .. 93,822.36 Postage * Me 228.38 Balance met by Society .. ms 600.54 $4,732.90 $4,732.90 $ Net cost Proceedings oy ee BP} 655.54 Net cost Reprints .. a ae fe 286.52 Total net Publication Costs be $942.06 The accounts have been set out in detail this year so as to show the true net cost. In order to comply with Post Office regulations, a proportion of every member’s subscription must be set aside specifically as a subscription to the Proceedings ; Council decided that this should be $5.00 per annum. ‘‘ Sub- scriptions ”’ therefore include members and other subscribers, including the State Government, which purchases 100 volumes each year. Membership During the year 11 new members were admitted to the Society, three resigned, and four were removed from the list of members. The numerical strength of the Society at 1st March 1972, was: Ordinary Members, 267; Life Members, 26; Corresponding Members, 3; total, 296. Professor Sir Rutherford Robertson, K.B.E., C.M.G., F.A.A., F.R.S., was honoured by the Queen in the 1972 New Year’s Honours List with a knighthood. Monthly Meetings Addresses were given at the June and July meetings, and a Symposium was held at the October meeting; a Field Day excursion took the place of the PROCEEDINGS OF THE LINNEAN Society or New SoutH Watss, Vou. 97, Part 1 2 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR Ordinary Monthly Meeting in September, while Notes and Exhibits were a feature of the November meeting. Details of these may be found in the Abstracts of Proceedings in Part 4, Vol. 96 (1971). . Inbrary During the year it was decided to retain the library, and a part-time librarian was appointed. In order to make additional space available, most of the mono- graphs have been sent to Fisher Library, Sydney University, on permanent loan ; a suitable book-plate will be inserted in each volume by the University, indicating that it is from the Linnean Society’s collection. The serials have been re-shelved into alphabetical order as in the Scientific Serials catalogue; we hold 1,595 serials, of which 565 are current. A new catalogue is now in preparation, and it is hoped that when this is complete it will be possible to circulate a list of our holdings to interested libraries. Subsequently, it is proposed to catalogue those monographs, mainly from the British Museum, still remaining in the Society’s library. Linnean Macleay Fellowship Dr. A. Anne Warren (née Howie) completed her Fellowship in the Depart- ment of Geology, University of Melbourne. A paper entitled ‘‘ A Brachiopod Labyrinthodont from the Lower Trias of Queensland ”’, based on her research during the year, was published in Vo]. 96, Part 4 (1971). Mrs. Lynnette A. Moffat was appointed Linnean Macleay Fellow in the School of Biological Sciences, Sydney University, for one year as from the 1st July 1971. Her topic of research is the relationship of amphibians and reptiles, using recently developed methods such as cytotaxonomy and serology. Linnean Macleay Lectureship in Microbiology Dr. K. Y. Cho, Linnean Macleay Lecturer in Microbiology, University of Sydney, as from 1st January 1969, reported as follows on his work for the year to December, 1971: ‘‘ ‘Work has been completed on the investigation of aspects of the internal organization of Hscherichia colt by electron microscope studies of auxotrophs defective in envelope synthesis. Spermine treatment of a diaminopimelic acid-requiring strain under conditions of diaminopimelic acid starvation (thereby blocking cell envelope synthesis) resulted in the detection of a polar organelle. This organelle is connected to the plasma membrane and consists of ribosomes In association with a structure in interlocking electron dense and light com- ponents. It is proposed that this structure is an apparatus for the synthesis and assembly of protein and phospholipid components of the cell envelope, including the organization of cell division. If some of the electron-light components represent nucleic acid, then this organelle may also be involved in the replication and transfer of DNA. This is the first organeJle other than membrane invaguna- tions that has been discovered in Escherichia coli or any prokaryote. ‘““The work has been accepted for publication in the Australian Journal of Biological Science. ‘‘ Other work under investigation includes the isolation of phospholipid requiring auxotroph of Azotobacter for the study of membrane biogenesis, and the effect of spermine on Azotobacter.” Finance The balance sheets were presented by the Honorary Treasurer, who pointed out that during the year the Society had been faced with unusually heavy expenditure, some of it of a non-recurrent nature, which had resulted in a deficiency of $5,253.07 in the General Account. PROCEEDINGS OF THE LINNEAN SocreTY oF NEw SourH WaAteEs, Vou. 97, Part 1 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 3 This had been due: (a) To the general inflationary increase in costs and services of all kinds ; (b) to the general increase in rates of salaries, which the Society is obliged to meet in order to obtain and keep suitable staff ; (c) to the retirement of our previous Assistant Secretary and obligation to provide long service leave and an appropriate pension ; (d) to the reorganization of our record and accounting systems and general office procedures, necessitating also the purchase of some new office equipment. (e) to the reorganization of the library, which involved the part-time employment of a trained librarian and an assistant. The costs of the last mentioned will be covered in part by monies received under the compensation claim from the Sydney Cove Re-development Authority. The accounts have been presented in a slightly different form this year, on the recommendation of our Auditor, in order to bring them more closely into line with modern business practice. The Bacteriology and Fellowships Accounts remain in a satisfactory condition. A new account has been opened, styled the Linnean Society of N.S.W. Scientific Research Fund. Monetary donations of $2.00 or over to this fund are exempt from income tax. Its funds can be used only for scientific research, but permission to capitalize donations to it and allow it to accumulate has been obtained. The Honorary Treasurer drew the particular attention of members to this fund and its taxation benefits and expressed the hope that, as and when they were able, they might contribute to it as a means of stimulating scientific research in the future in the tradition established by Sir William Macleay and continued thereafter by the Society. Report on Science Centre With support from the N.S.W. State Government in the nature of a Crown grant of what has become an increasingly valuable block of land, the Linnean and Royal Societies, together with the Institution of Engineers, were able to build Science House in 1930. Since then this building, on the corner of Gloucester and Essex Streets, has provided in Sydney a Science Centre with lecture hall, meeting room facilities, libraries and accommodation not only for the owner bodies, but also for tenants of the building and other scientific and educational groups, and often on a regular basis. The resumption of Science House in December 1970 led the Linnean and Royal Societies to request the Government of New South Wales to provide, as it had done in 1930, a new site for these Societies to re-house themselves. The Societies have resolved to accept responsibility for the building of a new Science Centre much as they had in the building of Science House some 40 years ago. It is expected that part of the new Centre will provide accommodation and Shared general facilities for professional and scientific organizations under favourable terms, while another part of the Centre will provide office space at commercial letting rates. The New South Wales Government, no doubt mindful of the history and contribution of both Societies within this State, has considered our request sympathetically, has encouraged us in our efforts, and has indicated that it will give as much support as possible to our scheme. Following the approval given by both the Linnean and Royal Societies for the building of a Science Centre, a Planning Committee was appointed, with representatives from both Societies. This committee has met regularly during the last year and has kept pace with the requirements, stage by stage, in the broad initial planning and in the clarification of the position of both Societies in relation to the project. Progress towards the establishment of a Science Centre may be Summarized under the following headings. PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, Vou. 97, Part 1 4 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR Compensation Science House provided for the Linnean and Royal Societies not only a home but also a source of income. The compensation which is being claimed from. the Sydney Cove Redevelopment Authority, the new owners of Science House following the resumption, is substantial. It is not yet known what the final amount will be, but this information should be available before long. It is the intention of both Societies to invest the compensation money, excluding the amount paid in respect to claims for disturbance, in the new Science Centre. Until the exact figure for compensation is known, and until the cost of erecting a building on a particular site has been calculated, it will not be possible to say how much additional finance will be required. However, the compensation money for the two Societies should provide enough cash in hand. An application has been made to the Sydney Cove Redevelopment Authority for advances to the two Societies on this compensation money, and we have been advised that cheques will be available subsequent to clearance from the Valuer-General’s Office. In short, the resumption of Science House will inevitably remove from the Societies their owned-home and source of income. The Societies, conscious of their long history and anticipated long future, have resolved to invest compensation money in such a way as to provide another owned-home and source of income. To do this, they are working jointly, knowing well that they will build a better Science Centre to fulfil their future needs if they combine their resources rather than act separately. Site and Tenure Two sites in the eastern Rocks Area, as well as other suitable sites, are under consideration. At a meeting with representatives of the two Societies on 9th March 1972, the Minister of Cultural Activities undertook to investigate the possibility of special concessions in the event of a site being chosen in the Rocks redevelopment area, where land leases may ordinarily be of rather limited duration. In the event of the Science Centre being erected in a suitable area, the Government would be prepared to undertake to rent a specified area of the income-providing office space of the Centre. Property Consultants and Legal Advisers Messrs. Jones, Lang, Wootton, property consultants to Science House and valuers for compensation purposes, were appointed by the two Societies to be Property Consultants for the Science Centre project. As property con- sultants, they will be responsible for a wide range of services which in the initial stages involve (a) general consultancy work leading to the establishment of a viable basis for development ; (b) negotiations in respect to a suitable parcel of land and the right to develop it to an agreed floor space ratio and for agreed uses ; (¢) negotiations for finance for the project. Later they will be involved in the co-ordination and direction of the development project, the leasing of the space created within the development surplus to the requirements of the Science Centre itself, and possibly the management of the completed development. Messrs. Stephen, Jacques & Stephen, Legal Advisers to Science House, have been appointed by the two Societies as legal advisers for the Science Centre project. Act of Incorporation The Act of Incorporation of the Linnean Society of New South Wales dates back to 1884, and under this Act, which has not been so far revised, the Linnean Society is far more restricted in its possible investments and hence sources of income than is the Royal Society of New South Wales ; it is also restricted in various other ways, even in respect to the Fellowships it offers. With the approval of its members, the Linnean Society is now in the process of amending PROCEEDINGS OF THE LINNEAN SociETy oF NEw SoutH WALES, Vou. 97, Part 1 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 5 its Act of Incorporation. The matter is at this stage in the hands of the Parlia- mentary Draughtsman and the Minister of Cultural Activities has offered his assistance in bringing forward an Enabling Act. Science Centre Company In order for the Linnean and Royal Societies to operate a Science Centre appropriately, it will be necessary for the Societies jointly to form a company. Preparation of a draft Memorandum and Articles of Association has been completed, and such a company could be established at short notice. Progress Reports In response to an early questionnaire, a considerable number of scientific and professional organizations expressed an interest in the possibility of accom- modation in the proposed Science Centre. These bodies are being advised from time to time of the stage reached in the project. A first progress report has been prepared and circulated. Government Approval of the Plan It is clear that the Government of New South Wales approves of our action in proceeding towards the establishment of a Science Centre. The Minister of Cultural Activities, the Director of the Sydney Cove Redevelopment Authority, Treasury officials and members of the Public Service Board have all given generously of their time and have made offers of help. It is realized by the two Societies, as also by the Government, that if the compensation payment reaches the level anticipated by the Societies the financing of the new Centre could proceed independently of the Government. However, if a new building on a new Site is to be a Science Centre in the real sense and is to provide low-cost facilities for other societies, then tangible assistance at least by way of concessions or guarantees will be required from the Government. We have been invited to discuss this matter further with the Minister of Cultural Activities at a later date. During the coming year it is hoped to see the finalizing of the financial aspects of the project, the appointment of an architect, and the completion of the architect’s brief outlining of the concept of the proposed Centre. Presidential Address The President then gave the Presidential Address, ‘‘ Evolution and Classi- fication in Hucalyptus ”’. Annual Elections No nominations of other candidates having been received, the Chairman declared the following elections for the ensuing year to be duly made: President: Dr. H. G. Cogger. Members of Council: Professor Donald T. Anderson, Professor Derek Anderson, Dr. L. A. 8. Johnson, Dr. P. J. Stanbury, Professor N. G. Stephenson, Dr. F. H. Talbot. Auditor: W. Sinclair & Co. The Chairman announced with great regret that Dr. A. B. Walkom had informed Council that he would not be able to nominate for Council membership again due to personal circumstances. As a token of its appreciation Council had appointed him Councillor Emeritus for the rest of his life. The meeting passed a resolution expressing appreciation for Dr. Walkom’s long and devoted service to the Society. The Chairman then installed Dr. H. G. Cogger as President. A vote of thanks to the retiring President, Dr. L. A. 8. Johnson, was moved. by Dr. Vickery and carried by acclamation. 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A. 8S. JOHNSON National Herbarium of New South Wales, Royal Botanic Gardens, Sydney [Delivered 29th March 1972] Synopsis Following general discussion of some principles of classification, some probable evolutionary trends in Hucalyptus are discussed, especially in relation to the recent classification of Pryor and Johnson (1971). Particular stress is laid on the multiple trends and varied final conditions in the calyx and corolla, which are more or less opercular, and in the inflorescence. Characters of anthers, ovules and seeds, cotyledons, hairs, oil glands and ducts, and associated insects are also reviewed as expressed in the eight subgenera recognized (including Angophora). Suggestions are made as to the phylogenetic connections of some subgenera and sections, and some problems of relationship are indicated within the eucalypts and between them and other Myrtaceae, which may respond to further detailed investigation and critical synthetic and analytical evaluation. A few changes in the Pryor and Johnson scheme are made or suggested. ‘‘ Our reasonings grasp at straws for premises and float on gossamers for deductions.” A. N. WHITEHEAD, Adventures of Ideas (1933). On CLASSIFICATION Four years ago (Johnson, 1968) I expatiated from this chair on the philosophy, methodology, problems, and especially the limitations of taxonomy in general. I concluded that no optimal classification was definable, much less attainable, whether on a phenetic or a phyletic basis, but that ‘‘ none the less, the elucidation of phylogeny can still proceed, and our unperfectable classifications can still be improved by reducing inconsistency until uncertainty or instability renders further change unprofitable’. No group of Australian plants has had so many investigators as the so-called genus Hucalyptus, yet such is its complexity and so many the gaps in our knowledge that its classification is still manifestly improv- able, and its detailed phylogeny still not elucidated. Recently my colleague Lindsay Pryor and I have published, in skeletal form, a new scheme of classification of the eucalypts (Pryor and Johnson, 1971). In deriving this scheme and in our own current revision of it, we have had to consider characters from as many fields as possible, and to evaluate them as to evolutionary significance. As I hope to have shown in 1968, a truly theory-free classification of organisms is an impossibility, although one may certainly develop classifications for which there is no defined theoretical foundation and in which the underlying theoretical assumptions are confused and frequently unconscious. Tf the distortions due to these assumptions are so multifarious and chaotic as to be evenly spread, constituting ‘‘ white noise’’, and if this background ‘‘ noise ’’ is not too great, some meaningful set of signals may be extracted from a phenetic analysis based on many characters. It is then up to us to interpret, in some scientifically or pragmatically profitable way, the results of the analysis. For this purpose, the ‘‘ results’ should not be taken as merely one particular hierarchical classification as represented by the dendrogram produced by a particular strategy. Whenever we proceed from the data to a dendrogram we lose information, and many topologically as weil as metrically different dendro- grams (and thus formal classifications) can be produced from a given set of data, merely by quite defensible manipulation of the strategies employed, as Lance. and Williams (1967 ; Williams, unpub. 1971) have very clearly shown. PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VotL. 97, Part 1 12 EVOLUTION AND CLASSIFICATION IN HUCALYPTUS If, then, we wish to make effective use of phenetic analysis to generate hypotheses (i.e. for “‘ interpretation ”’ of taxonomic data) we shall need to consider various dendrograms and also to check back frequently to the characters them- selves. Such hypotheses may relate to the prediction of properties (including genetic and physiological behaviour) of taxa and individuals. 30- flowered); it is the most usual condition in Hudesmia (umbellasters most commonly 3-flowered but in some species 7- or many-flowered): it is very widespread in Symphyomyrtus (umbellasters very often with 7, quite often with 3 or 11, Jess frequently with 1 or +15 flowers). Superposed twin umbellasters may be found in some axils, for instance not uncommonly in section SI Bisectaria ; these do not show any common rhachis or secondary bracts (so far as I know) and are probably a phylogenetically secondary development associated with the presence of supernumerary axillary buds. The S, condition of the curious ‘ Yellow Tingle”, SDA:A EH. guilfoylet (the sole species of section SD Tingleria) is fairly clearly due simply to reduction of subtending leaves to a bract-like state. Its continuing growth shows it to be basically distinct from the superficially similar ‘“‘ racemes of umbels ”’ of, say, JAA:A E. cloéziana (type T,) and to a lesser degree from the conflorescence of SWA:A H. microcorys (type §,;, elongated). Hudesmia, as mentioned, shows some odd conditions and the apparent ‘‘ umbels” in, for example, EHAACM EH. jucunda or HAADA E. gongylocarpa are probably clusters of umbellasters of type S,—but they need further investigation. In Symphyomyrtus, subtype 8S, occurs only once, in SNI:A LH. michaeliana. This rather perplexing species seems to be closest in leaf-venation, seeds, etc., to the ‘‘ Red Gums ” of section SN Hasertaria and its peculiar axillary triads of umbellasters are here tentatively interpreted as a secondary expansion of the 8, condition by an interpolated dichasial branching. A study of inflorescences in hybrids, if they can be made, may help to elucidate this. H. michaeliana needs more study morphologically, chemically, and by breeding experiments to determine whether our hypothesis as to its position is justified. It does not PROCEEDINGS OF THE LINNEAN SociETY or NEw SoutH WaAtgEsS, Vou. 97, Part 1 L. A. S. JOHNSON 23 seem to have much in common with SB Hquatoria and SS Howittaria to support any suggestion that its three-umbellaster groups are reduced T-type conflorescences. In section SU Adnataria, the ‘‘ true Boxes and Tronbarks”’, sub-type 8, is found as well as conditions transitional to, and fully characteristic of, sub-types S, and S,. These are the ‘‘ terminal, paniculate inflorescences’ of the older describers of the species concerned. The condition like T, and ©,, yields a massing of flowers towards the ends of the branches and doubtless has adaptive significance. Its derivation from 8, is clear, as is that of section SU from the basic stock of Symphyomyrtus. The ‘‘ Tallow-wood”’, SWA: A EH. microcorys (constituting section SW Sebaria) has a similar inflorescence but is an isolated species whose links with the rest of Symphyomyrtus are still obscure. The remaining features to be mentioned require, or allow, only a briefer treatment. References are given only when they are not covered in Pryor and Johnson (1971) or otherwise need special mention. (3) Androecium. (a) Anthers. Figures 4 and 5 summarize the anther types, which are more fully illustrated by Blakely (1934, 1965). The attachment of the filament to the connective is not shown, but the primitive dorsal attachment and versatile condition is indicated by the letter ‘‘v’’, and is retained in all groups except some sections of Symphyomyrtus. Here more basal attachment and the semi-versatile condition are associated with the series SIX Calycogonae (left anther of “sv” pair in Fig. 5) and SIZ Foecundae (right anther of ‘sv ” pair) of section SI Bisectaria, while completely adnate anthers with more or less pore-like openings are characteristic of section SU Adnataria. The usual (‘‘ porantheroid ’’) condition in the latter is as in the left-hand of the ‘‘a”’ pair, but in the three series SUT Polyanthemae, SUV Paniculatae, and SUX Mel- liodorae the so-called ‘‘ terminales-type ”’ anther (right-hand of the ‘‘a”’ pair) is found. No breeding barrier exists between species with these two anther types, and the division on anthers cuts across that on persistence of calycine operculum [see above under operculum-type C(A)S(U)]. Hence until more information is available on other characters we cannot say whether the anther or the operculum condition (or both) arose more than once. The generalized ‘‘ macrantherous’’, versatile anthers with long, separate dehiscence-slits are variously shortened in some series but these variants are not separately shown. Small anthers with short, somewhat divergent loculi (top left of Symphyomytus group) are found in SB and SS, the Hquatoria-Howittaria group, which may need to be excluded from Symphyomyrtus. A unique, specialized anther type (bottom ‘“‘v” of Symphyomyrtus group) characterizes the single species of SD Tingleria. The ‘‘ renantherous ”’ type with the loculi confluent at the top was once thought to be a defining character for the ‘* Renantherae’’. Krom these the sections SB and SW have now been removed, and it has been shown that the bulk of the ‘‘ Renantherae”’ belong together with a few species (series MAA Preissianae) which exhibit the primitive ‘‘ macran- therous ”’ anther type, and also with some intermediates (series MAB Diversifoliae) in Monocalyptus (in which PJ, recognizes only a single large section MA Renantheria). Thus the renantherous anther type is of later origin than the separation of the Monocalyptus line. (b) Other androecial features. Other features of the androecium are also useful in classification and of evolutionary interest but can only be mentioned. They include (i) the so-called staminophore (androphore would be a happier term) or stamina] ring, (ii) the grouping of stamens into four clusters evident in many Eudesmia species and occasionally elsewhere, e.g. in SW Sebaria, (iii) the development or otherwise of oil-glands in the filaments [a nice series is shown, for instance, from MAA Preissianae (abundant, large glands) through MAB PROCEEDINGS OF THE LINNEAN SooretTy oF NEw SoutH WaAtzsS, Vou. 97, Part 1 24 EVOLUTION AND CLASSIFICATION IN EUCALYPTUS Diversifoliae (few, small glands) to other series of Monocalyptus (without glands) |, (iv) the more or less staminodial condition of the outer stamens in certain groups, e.g. EAAD Odontocarpinae, SIX Calycogonae, SUJ Ochrophloiae, SUV Paniculatae, and SUX Melliodorae, a feature which has clearly had multiple origins, (Vv) various conditions in the flexure of filaments, which are inflexed in young buds even if straight in mature buds with long opercula such as those of SICB Cornutinae, and in the apposition of anthers to the disc* (e.g. in SL Dumaria). The detailed study of pollen by electron microscopy may yield information of classificatory and possibly phylogenetic value. Pollen-morphological examina- tion with the light microscope has not been very helpful in Hucalyptus. (4) Gunoecium and Associated Features. (a) Ovary, style, ete. No discussion of these features is yet possible which would be of much taxonomic or phylo- genetic significance, except as treated under (b). We may look hopefully to others currently studying the development and morphology to bring forth some illuminating discoveries in this field. (b) Ovule and seed. Arrangement of ovules and ovulodes on the placenta has been studied by Carr and Carr (1962, 1963) and may prove to have consider- able value in elucidating phylogenetic relations within Hucalyptus, and also between particular groups of Hucalyptus and other genera, as mentioned below under the discussion of possible polyphylesis. I have not yet examined placenta- tion and ovule arrangement extensively and cannot comment upon them in detail. Two basic ovule types, the anatropous and the hemitropous, are found in the eucalypts (Gauba and Pryor, refs. in Pryor and Johnson, 1971) and are associated with certain seed characters, especially the presence of a raphe in the anatropous type, as one would expect. These and other seed characters, notably the varying degree of development of the inner epidermis of the outer integument as a crystal epithelium, cannot be elaborated here. They strongly support the PJ, scheme, especially in the complete separation of Gaubaea from Hudesmia, the inclusion of EFC Miniatae in Hudesmia, the affinity of Angophora, Blakella, and Corymbia, and the marked difference between those three and both Hudesmia and Gaubaea. Sectional groupings within Symphyomyrtus are also supported by seed characters but these need more study. Ovule and seed types appear to associate Idiogenes with Gaubaea rather than with SB Hquatoria and SS Howittaria, although Idiogenes resembles these latter in its T-type conflorescence and Type ZS(B) operculum. Further comparisons may tell us in which of these characters convergence must be invoked. Figures 4 and 5 show only the anatropous-hemitropous distinction ; the anatropous is most general in Myrtaceae but campylotropous or hemitropous ovules do occur elsewhere in the family, for instance in Arillastrum (Dawson, 1970, and see below). Most interestingly, the seeds of GAA: A #. curtisii are extremely similar exteinally and anatomically to those of Tristania conferta. Tristania as currently recognized is a heterogeneous group and should probably be split into three genera. Clearly, the resemblances and differences between Gaubaea (and other subgenera) and the components of Tristania and their allies will need much deeper investigation. It is of course not necessarily the case that either all the hemitropous or all the anatropous subgenera belong together phylogenetically. (c) Fruit (including fruiting hypanthium). Features of the fruits are of course used a great deal in distinguishing and circumscribing the species and subspecies of Hucalyptus, and many of these features run through series, sections, or even subgenera. It is easy, for instance, to recognize an Angophora, a Blakella, * Carr and Carr prefer to term this the nectary ; it is nevertheless a floral disc in the general sense in which that term is employed in taxonomic description. PROCEEDINGS OF THE LINNEAN SocteTY oF NEw Soutu WaAtEs, Vou. 97, Part 1 L. A. 8S. JOHNSON 25 or a Corymbia by its fruit and, on a lower level, the fruits of Monocalyptus series MAH Capitellatae (the ‘‘ true Stringybarks’’) are all patently variations on a theme, as are those of Symphyomyrtus series SIC Cornutae or SIT Oleosae. It has not as yet been so easy to describe or define the whole congeries of characters which hold the major groups together, or to discern trends which may be phylo- genetically interpreted. (5) Cotyledons. These will not be discussed in detail. Some idea of the types characteristic of the subgenera is given in Figures 4 and 5. The ango- phoroid group A, B, C clearly hangs together and so do the sections of Symphyo- myrtus with their emarginate (bilobed) cotyledons (though the dubiously included SB and SS need to be checked). The right-hand figure for Symphyomyrtus shows the ‘‘ bisected’, Y-shaped cotyledons characteristic of the large section SI Bisectaria for which other positively uniting characters are stil) being sought, though the process of taxonomic agglomerative clustering (on a non-numerical basis at present) tends to associate the various series of this section. Y-shaped cotyledons occur also in SUUBA #. dundasii, which on other features appears to belong to section SL Dumaria, a group which seems to have radiated in parallel with Bisectaria but has retained the broader, bilobed cotyledons of the rest of Symphyomyrtus. The description by Carr and Carr of certain small outgrowths as cotyledonary stipules is open to further study and interpretation. Stipules are not a general feature of Myrtaceae. (6) EXmergences. (a) Trichomes (“hairs”). Hairs are found on various organs, most often juvenile shoots and leaves but also sometimes on adult shoots and even inflorescences and hypanthia. They need detailed anatomical study and their significance cannot be fully assessed. Type ‘‘a” (Figs 4, 5) comprises “‘ angophoroid ”’ hairs arising singly from undifferentiated parts of the epidermis. They are either uniseriately several- celled as in Angophora, or single-celled and very short as in Corymbia. Ihave not found hairs of this type in Blakella. Type ‘Sr?’ comprises radiating uniseriate hairs which arise in clusters from more or less raised oil glands. Three sub-types are recognizable at sight but not yet accurately described or analysed; I suspect that they have arisen inde- pendently in the course of phylogeny. They are found as follows: (i) subtype ‘*7(B) ” (blunt-ended, 1- or 2-celled, rather thin-walled hairs, often almost erect, on prominent glands or bristle-glands) in some species of Blakella, e.g. BAA: H H. gilbertensis. (ii) Subtype ‘‘r(E)”’ (blunt-ended, of rather thin-walled cells, on slightly prominent or flat glands) on juvenile shoots of all species of Hudesmia. (111) Subtype ‘‘r(M)”’ (acute-ended, of rather thick-walled cells, on prominent glands) on juvenile shoots of all species of series MAH Capzitellatae (these are the so-called ‘‘ stellate hairs’ of the Stringybarks) and of an undescribed species from south-eastern New South Wales (discovered after publication of PJ,) which may require the establishment of a new series or subseries in Monocalyptus. Somewhat papilliferous raised glands are found on a few species (e.g. MAKCA EH. regnans) of series MAK Obliquae, and these may be regarded as morphologic ally transitional between the ‘‘r(M)”’ condition and the hairless state of most of Monocalyptus. Hairs are reported from a few species of Symphyomyrtus ; the report by Maiden for SUABB HE. leptophleba is false and evidently due to mixture of material, that for SICBE L. lehmannii is based on blunt several-celled papillae on the surfaces of glands. Significantly, hairs are absent in Gaubaea and Idiogenes, which were referred to Eudesmia (as ‘‘ Hudesmieae”’) by Carr and Carr as recently as 1970 (see Pryor and Johnson, 1971 for discussion). Trichomes closely resembling the various eucalypt types do not seem to occur in other genera of the Myrtaceae, but simple hairs of rather different types are quite common in many genera. PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wates, Vou. 97. Part 1 26 EVOLUTION AND CLASSIFICATION IN EUCALYPTUS (b) Bristle-glands. These (Figs 4, 5) are characteristic of the angophoroid trio Angophora, Blakella, and Corymbia, and are essentially elongated protruding oil glands, with a multicellular wall. In Blakella they are very thin and may superficially resemble simple trichomes, which has led to misdescription of species of this group. Some degree of elongation of raised oil glands occurs in other Species and in young seedlings of MAKAA LH. obliqua, for instance, these are quite bristle-ike, though doubtless representing a parallel or convergent development. H. obliqua is quite unlike the angophoroids in other respects. (7) Ow Ducts and Glands in the Pith. These are described by Carr and Carr (refs. in Pryor and Johnson, 1971) and are distributed as shown in Figs 4 and 5 (information partly from Carr and Carr, partly new observation). Once again the angophoroid A, B, C are linked, by the possession of oil ducts in the pith. Presence or absence of glands may be a fairly regular feature in some sections or Series (e.g. most of the first half of the series of SI in the PJ, arrangement have pith glands, whereas SIP-SIZ inclusive lack them ; most of SL have pith glands ; SE, SN and SP lack them) but in other cases there is considerable variation even within series (as in those of SU). They are absent altogether from Gaubaea, Idiogenes, Monocalyptus, and SB Hquatoria and SS Howittaria. InSP Maidenaria oil glands are reported in the secondary bark of older trees of most species but not quite all; this feature appears to be peculiar to the section. It is difficult to interpret these conditions in terms of trends (or adaptations) but they do aid in a polythetic classification. It should be remembered that oil glands are almost always present in the primary cortex and in leaves and some floral parts. (8) Chemical Features. (References will be found in Pryor and Johnson, 1971, and in works cited therein). (a) Terpenes and other essential oil constituents. This is a classical field in eucalypt chemotaxonomy, and indeed the work of R. T. Baker and H. G. Smith and their successors is classical in chemotaxonomy generally. The patterns tend to support PJ, but caution is necessary in interpretation, due to the possi- bility of switches in metabolic pathways which may be determined by quite simple genetic changes, and not necessarily very rigidly stabilized by selection so far as the oil constituent end-products are concerned. Information is hoped for soon on such critical groups as Gaubaea, Idiogenes, SB EHquatoria and SS Howittaria. (b) Polyphenols. The information in this field, due chiefly to the work of Hillis, has been presented according to Blakely’s classification, and in a few critical cases is based on material of doubtful identification. I hope chemists will review it in relation to PJ, to see what light is shed thereby, and that they may be able also to interpret the significance of occurrences more clearly in the light of increased knowledge of biochemical pathways. At present I can only say that PJ, appears to be supported in general, especially in relation to the delimitation of Monocalyptus which contains (though not invariably) the distinctive substance renantherin. (c) Leaf waxes (chemistry and shape of cuticular deposits). As pointed out by Pryor and Johnson (1971) this has proved a disappointing field, despite the thorough studies by Hallam and Chambers. Although there is a general fit to the broad classification of PJ,, there is evidence of much parallelism, and the Shapes revealed in the electron-micrographs seem often to be associated simply with glaucous versus non-glaucous conditions as seen with the naked eye, different types being found in obviously closely related species which are often within a Single superspecies. PROCEEDINGS OF THE LINNEAN SocieTy oF New SoutH Wates, Vou. 97, Part 1 L. A. 8S. JOHNSON 27 (d) As reflected by host-insect associations. This field has not been at all thoroughly surveyed but the large, lerp-forming genus Glycaspis (Hemiptera : Psyllidae) has been intensively studied taxonomically by Moore, and its associa- tions are interesting at the subgenus level. No Glycaspis at all is known from Angophora,* Blakella or Corymbia, although one subgenus, Boreioglycaspis, is found on Melaleuca, which is only rather remotely related to any eucalypt. Gaubaea and Idiogenes are insufficiently examined but no Glycaspis is recorded from those subgenera either. The type subgenus Glycaspis occurs widely on Symphyomyrtus (no information from SB-SS, SD, and no record from SW although this has been searched) and on a few species of Hudesmia (both EA and EF) aS well as, oddly enough, on T'ristania conferta (which does not seem to be related to these eucalypt groups—see above). On the other hand the numerous species of subgenus Synglycaspis are confined to Monocalyptus. At sectional and lower levels in the eucalypts the occurrences of Glycaspis species seem to me to have much less taxonomic significance, and I would certainly not agree with Moore’s suggestions that eucalypt-Glycaspis associations throw doubt on eucalypt relationships which are firmly established on other grounds. One can hardly do this when one finds the same Glycaspis species on species of SI and SU (e.g. G. repentina) or even of EA and SN (on both of which G. onychis occurs). One could go on to discuss other features, e.g. wood and bark anatomy, leaf venation patterns, fungal pathogen susceptibility (such as the association of Phytophthora cinnamomi and Monocalyptus, or Ramularia and the angophoroids), epidermal anatomy (a possibly promising field in which work has begun by scanning electron microscope techniques by the Carr group and others). But this would not add much to the picture at this stage. Neither would the scanty fossil evidence, which provides no detailed background of the phylogenetic history. Chromosome numbers are depressingly constant (see Pryor and Johnson, 1971). DISTRIBUTION OF GROUPS AND CHANGES IN THE CLASSIFICATION Distributions of the subgenera and sections are of interest and are shown in a general way in Table 1. It would perhaps be better to use phytogeographic divisions rather than States, but to do so would have involved difficulties of compilation. Table 1 is compiled to recognize some small changes from PJ,; we know a little more about some distributions, and with further study have altered our views on a few species and subspecies. Apart from the species mentioned above under ‘‘ Trichomes ’’, the most interesting specific case concerns EH. pachycalyz, referred with doubt in PJ, to the synonymy of SNABAA E. alba [ssp. alba], in the absence of the Type specimen (apparently lost while on loan some twenty years ago) or any other material. H. pachycalyxr has now been rediscovered and, having examined twigs, leaves, buds, flowers, fruits, seeds and cotyledons, I am convinced that it belongs in section SI Bisectaria. I here assign it the code SIQ:E, placing it in series SIQ Squamosae, although it differs in bark type and other details from SIQ:A H. squamosa, the other strictly eastern member of Bisectaria [SIR:E HE. bakert is merely the eastern vicariant of the central and western SIR:A ZH. jutsonii s. lat.]. It is most remarkable to find a true Bisectaria in north-eastern Queensland, adding to the mystery of why this species and H. squamosa should have reached the humid east. In other series of Bisectaria only eremaean species appear to have crossed the continent. Likewise, why did Bisectaria radiate and flourish so exceedingly in the west while its Squamosae representatives (which are hardly primitive in the section) barely hang on in the east ? Another change from PJ, is the elimination of section SQ Umbrawarria. T am now convinced that its sole species, H. umbrawarrensis, is closely related to * Except one very doubtful record of subgenus Glycaspis on BAA:A E. tessellaris. PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wates, Vou. 97, Part 1 28 EVOLUTION AND CLASSIFICATION IN EUCALYPTUS SNABG H. brevifolia, and assign it the new coding SNABI. Further, ‘“ SLI:G E. comitae-vallis ” is in fact, so far as the Type specimen is concerned, a probable hybrid: SLI:I LZ. concinna x SUUAA EL. ovularis (syn. EL. cylindrocarpa, not the un- described SLUAC to which the name ‘‘ H. ovularis”’ has been misapplied in Western Australia). H. brachycorys is not, then, a subspecies of “‘ SLI:G” but appears to be a member of series SLU Dundasianae subseries SLUA Ovularinae, and I here assign it to the coding SLUUAK (Mr. M. I. H. Brooker, pers. comm., suggested this affinity for H#. brachycorys, and I agree with it). A question for the future will be to determine whether there are general adaptive tendencies in the several sections and subgenera which are linked to their present distributions and the climates and conditions of their geographic origins, if those can be determined. ARE THE EUCALYPTS POLYPHYLETIC ? Elevation of some or all of the subgenera to generic status would be obligatory, at least in my view, if we were to become convinced that their phyletic relationships lay more closely with other genera of the Myrtaceae than with each other. I suspect that this may in fact be so. It is possible, as we have seen, that Gaubaea is related to Tristania, while Angophora-Blakella-Corymbia may well be more closely related to Arillastrum (Spermolepis), a very interesting New Caledonian genus studied by Dawson (1970). Dawson has shown that Arillastrwm should not be placed in Metrosiderinae if that subtribe is reasonably restricted in its content and circumscription. Unlike the Metrosideros alliance, Arillastrum has stamens (some staminodial) in many whorls, ovulodes as well as normal ovules and these arranged in a definite pattern, campylotropous ovules (not very different from the ‘‘ hemitropous”’ type of some eucalypts), a crystal layer in the seed-coat, and broad reflexed cotyledons. The petals are not as broad-based as in Angophora but are less clawed than in many Myrtaceae and have a median thickened area. I emphatically do not suggest that Arillastrum is to be regarded as representing any kind of ancestor of any group of eucalypts ; nevertheless it shares a number of characters with several of the ‘‘ hemitropous ” set of eucalypt subgenera, although it differs variously from all of them. Dawson has pointed out that Tristania (s. lat.), and Xanthostemon also, should be separated from the Metrosideros group, although they have retained the anatropous ovule condition. Hucalyptopsis, which resembles eucalypts in seeds, cotyledons, and of course in having an operculum, may also be related to some of the eucalypt lines. Correlation of the results of Dawson’s continuing work with that of students of ‘‘ Hucalyptus’’ (I dare now to put it in quotes!) should be of the greatest interest to the phyletic taxonomist and to the phytogeographer. We must, I think, seek the origins of the eucalypt lines near the very roots of subfamily divergence in the Myrtaceae, although I would not support the suggestion of Pilipenko (1962) that ‘‘ Hucalyptus ” is derived from “‘ Hugenia ” (itself a hetero- geneous assemblage)! If the eucalypts are indeed polyphyletic in this broad (subtribal) sense, then they present an even more remarkable example of parallel and convergent evolution, presumably in response to environmental selection, than has been thought. CONCLUSION I have discussed with varying degrees of sketchiness some of the observed characters and the apparent trends in them, in relation to the PJ, classification. Naturally I consider that they support this classification, within the general limitations of classifications as discussed at the beginning of this address. If I did not, then I should change the classification, and have indeed suggested some possible changes for checking. The suggestion of polyphylesis from rather widely separate origins is, I suppose, the most radical and far-reaching, but it PROCEEDINGS OF THE LINNEAN SocrETy or New South WALES, Vou. 97, Part 1° L. A. 8. JOHNSON 29 calls for more substantial evidence before incorporation into the formal system. Uncertainty at this level does not vitiate the detailed system at lower levels. Space is lacking here to recapitulate the various and complex hypotheses involved ; they are implicit in PJ,, to some extent in Fig. 1, and in the foregoing discussion—and they can be dragged into the light by those who will, I hope, help to investigate them. As a gratuity to those critics who regard speculation as a scientific sin, I refer them to the quotation from A. N. Whitehead which is the ironic text for this address. Iam not very attracted by formalism and rigidity in science (though I am grateful that there are some who are) and would venture that even in the Queen of the Sciences mathematicians of insight and wide-ranging interest would agree that somewhat imprecise visualization of things as a whole, and gathering of many threads of thought and fact, can be as profitable as the equally necessary punctilious following-through of detail. In this small field of eucalyptology, such a two-pronged approach should also be productive. We have generated some hypotheses—it is for the next few years to show how good or bad they are. ACKNOWLEDGEMENTS I thank Professor L. D. Pryor, Mr. D. F. Blaxell, and Mr. M. I. H. Brooker for providing information; Dr. Barbara G. Briggs read and constructively criticized the manuscript, and [ am also grateful to her as well as to Mrs. Jennifer Walsh for assistance with the figures. References Buake, 8. T., 1953.—Studies on northern Australian species of Eucalyptus. Aust. J. Bot., 1: 185-352. Buiaxkety, W. F., 1934.—*‘ A Key to the EKucalypts.” Ist ed. (The Worker Trustees, Sydney.) , 1965.—*‘ A Key to the Eucalypts.” 3rd ed. (Commonwealth of Australia Forestry and Timber Bureau, Canberra.) Carr, D. J., and Carr, Stetyta G. M., 1959.—Developmental morphology of the floral organs of Hucalyptus. I. The inflorescence. Awst. J. Bot., 7: 109-141. , and , 1962.—Natural groups within the genus Hucalyptus. In “ The Evolution of Living Organisms.” (Symposium.) (Royal Society of Victoria, Melbourne.) Carr, STELLA G. M., and Carr, D. J., 1963.—The taxonomic position of certain eucalypts. Proc. Roy. Soc. Vict., 77: 207-216. Dawson, J. W., 1970.—Pacific capsular Myrtaceae. I. Reproductive morphology of Arillastrum gummiferum Panch. ex Baillon (New Caledonia). Bluwmea, 18: 431-440. GuHIsELIN, M. T., 1969.—The principles and concepts of systematic biology. In “* Systematic Biology ”, 45-55. (National Academy of Sciences, Washington.) Hott, D. L., 1967.—Certainty and circularity in evolutionary taxonomy. Evolution, 21: 174-189. Jounson, L. A. S., 1968.—Rainbow’s end: the quest for an optimal taxonomy. (Presidential address.) Proc. Linn. Soc. N.S.W., 93: 8-45. , 1970.—Rainbow’s end: the quest for an optimal taxonomy. Syst. Zool., 19: 203-239. (Reprint of Johnson, 1968, with addendum.) , and BLuaxety, D. F., in press a.mNew taxa and combinations in Hucalyptus—l. Contrib. N.S.W. natn. Herb., 4 (No. 5). , and ———__., in press b.—New taxa and combinations in Hucalyptus—II. Contrib. N.S.W. natn. Herb., 4 (No. 6). Lance, G. N., and Wittiams, W. T., 1967.—A general theory of classificatory sorting strategies. I. Mierarchical systems. Computer J., 9: 373-380. PiuiPenko, F. S8., 1962.—Proiskhozhdenie i évoliutsiia roda évkalipt. (The origin and evolution of the genus Hucalyptus). Bot. Zhurn., 47: 188-201. Pryor, L. D., and Jounson, L. A. S., 1971.—‘‘ A Classification of the Eucalypts.” (Australian National University, Canberra.) , and Knox, R. B., 1971.—Operculum development and evolution in eucalypts. Aust. J. Bot., 19: 143-172. Trou, W., 1964.—*‘ Die Infloreszenzen ”’, 2, pt. 1. (Fischer, Stuttgart.) Waker, J., and Bertus, A. L., 1971.—Shoot blight of Hucalyptus spp. caused by an undescribed species of Ramularia. Proc. Linn. Soc. N.S.W., 96: 108-115. Wittiams, W. T., unpub. 1971.—Principles of clustering. (MS.) Note: For other authors mentioned in the text without dates the references (sometimes numerous) will be found in Pryor and Johnson, 1971. PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 97, Part 1 A BROODING ECHINOID FROM TASMANTA ALAN J. DARTNALL The Tasmanian Museum, Hobart, Tasmania [Accepted for publication 15th September 1971] Synopsis A brooding species of echinoid which belongs to the genus Pachycentrotus is described. It is of interest because only one other non-cidaroid, regular urchin is known to brood its young. The species is also endemic to the Maugean marine fauna of south-eastern Tasmania. INTRODUCTION The sea urchin genus Pachycentrotus was erected by H. L. Clark (1912) to accommodate Sphaerechinus australiae A. Agassiz. The genus is restricted to the waters of south and south-eastern Australia and, hitherto, was considered to contain only the one species. A further species considered to belong to the genus is described below. : Family STRONGYLOCENTROTIDAE Genus Pachycentrotus H. L. Clark, 1912 Pachycentrotus bajulus sp. nov. Description of Holotype A small echinoid with a slightly depressed test ; hd.=33 mm., vd.=21 mm. The diameter of the apical system is 10-6°%, hd. and the diameter of the peristome is 30% hd. Viewed from above the test looks like a rounded pentagon because the ambulacral areas are slightly inflated and project beyond the level of the interambulacra. There are 22-23 plates in each column of the ambulacra. Each plate carries a large, centrally placed primary tubercle which increases in size from the apical system to about the eighteenth plate and then becomes smaller as the ambulacra approach the peristome. Two other tubercles are distinct features of the ambulacral plates (Fig. 1 (b)). Firstly, one tubercle is set on the lateral apex of the ambulacral edge of the plate, and secondly, a tubercle is situated beside and below the large primary tubercle and between that tubercle and the poriferous area of the plate. Other small tubercles are present on the ambulacral plates, two or three being present on the plates near the apical system and about sixteen on the largest ambulacral plates. Most of the ambulacral plates are perforated by four pairs of pores arranged in an arc of three pore-pairs on the outside edge with the extra pair placed towards and below the primary tubercle of each plate. Towards the periproct and peristome only three pairs of pores are present, two at the edge and oneinset. Onpreliminary examination it appears that the pore-pairs are arranged in ares of three or four because the inner pair of pores of the preceding plate is aligned with the outer pore-pairs of the following plate. There are 18-19 plates in each column of the interambulacra. Each inter- ambulacral plate carries a large primary tubercle towards the lower edge of the plate (Fig. 1 (a)). The arrangement of smaller primary and secondary tubercles is also shown in the figure. The tubercles on the test are all smooth and imperforate. The apical system of the holotype is illustrated in Fig. 2. The genital and ocular plates each carry a large primary tubercle similar to those found on the ambulacral and interambulacral plates. Ocular plates II and III are exsert, PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WALES, Vou. 97, Part 1 ALAN J. DARTNALL 31 Fig. 1. Pachycentrotus bajulus sp.nov. (a) Interambulacral plates 6-8 from apical system. (6) Ambulacral plates 6-8 from apical system. Fig. 2. Pachycentrotus bajulus sp.nov. Genital and ocular plates. the remainder insert. The madreporite occupies most of genital plate IIT and is inflated, projecting above the level of the other ocular plates. In the holotype the periproctal plates are lost. About a dozen, irregularly arranged, imbricate plates are present in a denuded specimen of 20mm. hd. The anus is situated to one side of the periproct. The peristomial membrane carries numerous small slightly imbricating plates and ten ovoid, perforate plates for passage of the buccal tube feet. Small open gill slits are present at the perimeter of the peristome. PROCEEDINGS OF THE LINNEAN Society oF NEw SoutTH WALES, Vou. 97, Part 1 32 BROODING ECHINOID FROM TASMANTA The primary spines are about 6 mm. in length, possess a milled ring at the base, and are longitudinally ridged. Most are about twice as broad above the milled ring as at the tip and commence to taper about halfway along the spine. Some of the primary spines are slightly expanded at the tip. The secondary spines are about 2mm. in length, are slightly thorny along the longitudinal ridges, and are more club-shaped than the primary spines. Large globiferous pedicellariae are common. Tridentate, ophiocephalous and two kinds of triphyllous pedicellariae are also present (see Fig. 3). Colour. The interambulacral plates and the imperforate parts of the ambulacral plates are pale green. The colour is most intense around the apical system and fades away until, close to the peristome, the test is the off-white colour common to the poriferous areas. The madreporite is buff coloured. The primary spines are variously banded with dull violet-brown and green and the tips of the spines are white. Type locality. Darlington Beach, Maria Island, off the east coast of Tasmania. Latitude 42° 44’S. Longitude 149° 05’ EH. Holotype. A dried, partially denuded specimen from the type locality. Taken at low tide under rocks, 15.iv.1968. Coll. A. J. Dartnall, Tasmanian Museum Reg. No. H603. Paratypes. One dried specimen with 17 young amongst the spines of the test (Tasmanian Museum Reg. No. H604), four spirit specimens (T.M. Reg. No. H605), and three spirit specimens (Australian Museum Reg. No. J7801). All taken from under stones at low tide. February, 1948, Eaglehawk Neck, 8.E. Tasmania. Coll. V. V. Hickman. One dry specimen from Eaglehawk Neck, Tasmania, 27.1.1928, coll. W. Irwin Smith (Australian Museum Reg. No. J6393). Three spirit-preserved specimens from under stones at low tide, Adventure Bay, Bruny Island, S.E. Tasmania ; 6—15.1.1937, coll. V. V. Hickman (National Museum of Victoria Reg. No. H155). Htymology. The specific name is derived from the Latin masculine noun bajulus, meaning porter, in reference to the brooding habit of the animal. Affinities. Pachycentrotus bajulus is most clearly related to Pachycentrotus australiae (Agassiz). I have not seen any specimens from Tasmania which I would refer to P. australiae. One of the Tasmanian specimens described by H. L. Clark (1938, p. 404) is certainly P. bajulus (this is paratype J6393), but I have not seen the remainder of the series. Comparative comments here rest on the information in the literature and two specimens of P. australiae from the collections of the South Australian Museum. P. bajulus is a smaller animal than P. australiae, maximum recorded hd. being 33 mm. and 38 mm. respectively. The gill clefts are deeper in P. australiae and the pair of curved processes which extend from the epiphyses of Aristotle’s lantern and retain the tooth, are longer and more pointed in P. bajulus. The interambulacral plates of P. australiae carry more spines than those of the new species and the large, primary boss characteristic of P. bajulus is not present in that species. The globiferous pedicellariae of P. bajulus have deeper shoulders with more acute corners than those of P. australiae and the palms of the triphyllous pedicellariae are more expanded in the new species. Remarks. Pachycentrotus bajulus is the only non-cidaroid, regular urchin from Australia known to brood its young. The only other echinoid in this category is Hypsiechinus coronatus (family Temnopleuridae), which is limited to the far northern North Atlantic (Hyman, 1955). In H. coronatus the young are PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, Vou. 97, Part 1 O:\S mrn 33 ALAN J. DARTNALL O-2 mm £ E cs) 2 fe) £ £ ) 9° co) SS e Fig. 3. Pachycentrotus bajulus sp.nov. (a) Jaw of globiferous pedicellaria. (b) Jaw of tridentate pedicellaria. (c) Jaw of small triphyllous pedicellaria. (d) Jaw of large triphyllous pedicellaria. (e) Jaw of ophiocephalous pedicellaria. PROCEEDINGS OF THE LINNEAN Society or New Souruw WateEs, Vou. 97, Part 1 34 BROODING ECHINOID FROM TASMANIA brooded around the plates of the periproct. In P. bajulus the young ride scattered amongst the spines over the whole test. There is no marsupium or preferred brooding area. The holotype carried 12 young, ranging from 1-5 mm. to 5:5 mm. hd., when it was collected. Paratype H604 carries 17 young ranging from 1-3 mm. hd. (illustrated in Australian Natural History, in press). In a zoogeographical context the new species is also of great interest because it shows some parallels with the distributional and reproductive phenomena already reported in some sea stars from Tasmania (Dartnall, 1970). In this case P. australiae is replaced by P. bajulus in south-eastern Tasmania. The reproductive cycles of both species are unknown, though P. australiae is said to have small eggs and, probably, pelagic larvae (Mortensen, 1943). However, reproductive isolation of the two species may be inferred. P. bajulus, so far as is known, is restricted to south-eastern Tasmania and is a further example of the endemic fraction of the Maugean fauna which characterizes the area (see Bennett and Pope, 1960, and Dartnall, in press). It is of interest to note here that recent work has shown that a chiton, Ischnochiton mayi, with a similar 8S.E. Tasmanian distribution, is also a brooding form (EK. Turner, personal communication). The occurrence of brooding animals in the shallow water fauna of the south of Tasmania when non-brooding closely related species are found in warmer, more northern waters may make it possible to elucidate some of the factors which make brooding and viviparity advantageous in a habitat. Also, in any consideration of the effects of pollution on the marine fauna of the area it must be emphasized that replacement of this part of the fauna from adjacent areas will be slow (Chia, 1970) or, if modification of the habitat is extensive, nearly impossible. ACKNOWLEDGEMENTS I thank Emeritus Professor V. V. Hickman for his generous donation of material. References BENNETT, I., and Pops, C. E., 1960.—Intertidal zonation of the exposed rocky shores of Tasmania and its relationship with the rest of Australia. Aust. J. Mar. Freshw. Res., 11: 182. Crark, H. L., 1912.—Hawaiian and other Pacific echini. Mem. Mus. Comp. Zool., 34: 205. , 1938.—Echinoderms from Australia. Mem. Mus. Comp. Zool., 55. Cura, F. S., 1970.—Reproduction of Arctic marine invertebrates. Marine Pollution Bulletin, 1 (N.S.), 5: 78-79. Dartnatu, A. J., 1970—The Asterinid sea stars of Tasmania. Proc. Roy. Soc. Tasm., 104: 73. , in press.—Littoral Biogeography, in Biogeography and Ecology in Tasmania. W. Williams ed. Monographie Biologicae, Junk, The Hague. Hyman, L. H., 1955.—The Invertebrates. Vol. IV, Echinodermata. McGraw-Hill, New York. Mortensen, Th., 1943.—Monograph of the Hchinoidea III (3). Camarodonta II. Echinidae, Strongylocentrotidae etc. Reitzel, Copenhagen. PROCEEDINGS OF THE LINNEAN SociETy oF NEw SourH WALES, Vou. 97, Part 1 THE OCCURRENCE OF ROOT NODULES IN THE GINKGOALES, TAXALES, AND CONIFERALES A. G. KHAN* AND P. G. VALDER School of Biological Sciences, The University of Sydney (Plate 1) [Accepted for publication 17th November 1971] Synopsis The roots of 57 species of the Ginkgoales, Taxales, and Coniferales were examined for the presence of nodules of the Podocarpus-type and for the occurrence of mycorrhizal associations. Of the conifers, those placed in the Araucariaceae, Podocarpaceae, and Sciadopityaceae all bore nodules, whereas species of the Cupressaceae, Pinaceae, and Taxodiaceae did not. No nodules were observed on Gingko biloba or Taxus baccata. Vesicular-arbuscular mycorrhizas were of general occurrence amongst the plants examined except for the members of the Pinaceae, which bore the ectotrophic type. INTRODUCTION Records of the occurrence of root nodules amongst the Ginkgoales, Taxales and Coniferales have been tabulated by Allen and Allen (1965) and a fuller report of the occurrence of structures variously described as exostoses, mamelons, tubercles, and nodules is given in Table 1. However, apart from the records for Podocarpus spp. by numerous authors, for Dacrydium franklin, Micro- cachrys tetragona, Phyllocladus trichomanoides and Saxegothaea conspicua by Sprat (1912), for Pherosphaera hookeriana (Microstrobus niphophilus) and P. fitegeraldui (M. fitegeraldii) by Saxton (1930a, 1930b) and for species of Agathis, Dacrydium, and Phyllocladus by Baylis et al. (1963), no evidence has been reported which indicates that the structures described differ in any way from short roots. TABLE | Records of the Presence of Nodules Amongst the Coniferales Family, Genus and Species Literature Citations ARAUCARIACEAE Agathis australis .. fs .. Cockayne, 1921; Yeates, 1924; Bieleski, 1959; Baylis et al., 1963; Morrison and English, 1967. A. robusta ae ss .. Janse, 1897 (as Dammara robusta). A. vitiensis Ate, By .. Allen and Allen, 1965. Araucaria spp. .. be .. Hooker, 1854. A. angustifolia .. Sf .. Daugherty, 1963. A. heterophylla .. ie .. Janse, 1897: Yeates, 1924 (both as A. eacelsa). CUPRESSACEAE Cupressus sp. a se .. Hooker, 1854. C. sempervirens .. ae .. Janse, 1897 (as C. fastigiata). Tnbocedrus bidwillii ae: .. Yeates, 1924. Sabina chinensis oe .. Janse, 1897 (as Juniperus chinensis). Thuja sp. (Platycladus ?) .. Hooker, 1854. PoDOCARPACEAE Acmopyle pancheri a .. Sahni (1920). Dacrydium sp... ee .. Hooker (1854). * Present address: University of Panjab, New Campus, Lahore, West Pakistan. PROCEEDINGS OF THE LINNEAN SocteTy or New SoutH WALES, Vou. 97, Part 1 ROOT NODULES IN GINKGOALES, TAXALES, CONIFERALES TABLE 1—Continued Records of the Presence of Nodules Amongst the Coniferales—Continued Family, Genus and Species Literature Citations PoDOCARPACEAE—Continued . bidwillaa . biforme . colensor . cupressinum . . intermedium . franklinia . kirk . laxifolium Microcachrys tetragona Microstrobos fitzgeraldi M. niphophilus Phyllocladus sp. P. alpinus Be P. glaucus ae P. trichomanoides Podocarpus spp. SISSISSS S's . acutifolius . blumet .. . dacrydioides elatus elongatus falcatus. . ferrugineus gracilior halla henkelvi imbricatus latifolius . macrophyllus . macrophyllus var. maki nage pial Gt aj wel asad gel pelos 3o Sy . neriifolius . nwalis nubigenus * prostrata ”’ known) rosprigliosii salignus spicatus spinulosus totara “ variegatus ” . wallichianus Saxegothaea conspicua (no such name (a cultivar 7) rity ry ytd hey tht TAXODIACEAE Cunninghamia sp. Taxodium sp. SCIADOPITYACEAE Sciadopitys verticillata Yeates, ex Allen and Yeates, 1924; Yeates, 1924. Allen, 1965. Baylis et al., 1963. Yeates, ex Allen and Allen, 1965; Baylis et al., 1963. Yeates, ex Allen and Allen, 1965: Baylis et al., 1963. Sprat, 1912. Yeates, ex Allen and Yeates, ex Allen and Sprat, 1912. Saxton, 1930a, 1930b Saxton, 1930a, 19306 Allen, 1965. Allen, 1965. (as Pherosphaera fitzgeraldv). (as Pherosphaera hookeriana). Hooker, 1854. Baylis et al., 1963. Yeates. ex Allen and Sprat, 1912. Hooker, 1854; von Tubeuf, 1896, according to Nobbe and Hiltner, 1899; Bond, 1959. Yeates, ex Allen and Allen, 1965. Becking, 1965. Allen, 1965. Hooker, 1865; Petri, 1903; Yeates, 1924: Baylis et al., 1963. Petri, 1903; McLuckie, 1923. Petri, 1903: Sprat, 1912; Phillips, 1932 (both as P. elongatus and P. thunbergu var. angustifolia). Phillips, 1932. Yeates, 1924 ; Parker, 1932. Yeates, 1924: Phillips, 1932. Janse, 1897 (as P. cupressinus). Saxton, 1930 (as P. thunbergiz) : 1967. Nobbe and Hiltner, 1899 ; Schaede, 1943 (all as P. chinensis) : Becking, 1965. von Tubeuf, 1896, according to Nobbe and Hiltner, 1899: Shibata, 1902 (both as P. nageza); Becking, 1965. van Tiegham, 1970, according to Becking, 1965; von Tubeuf, 1896, according to Shibata, 1902; Egle and Munding, 1951; Becking, 1965. Yeates, ex Allen and Allen, 1965: Bond, 1967. Schaede, 1943. Yeates, ex Allen and Allen, 1965. Baylis et al., 1963. Baylis eé al., 1963. Phillips, 1932; Bond, Shibata, 1902; Petri, 1903; Petri, 1903. Furman, 1964: Becking, 1965. Sprat, 1912; Bottomley, 1913 (both as P. chilina). Yeates, 1924; Baylis et al., 1963. MeLuckie, 1923. Sprat, 1912; Yeates, 1924; Baylis et al., 1963. Ferreira dos Santos, 1947, ex Allen and Allen, 1965. Petri, 1903 (as P. latifolia Wall.). Sprat, 1912. Hooker, 1854. Hooker, 1854. According to Hooker, the _ elder de Candolle had earlier noted exostoses on T’. distichum (Theorie Hlementaire, Ed. 2, p. 356). Uemura, 1964. PROCEEDINGS OF THE LINNEAN Society or NEw Soutru WaAtgss, Vou. 97, Part 1 A. G. KHAN AND P. G. VALDER 37 Allen and Allen (1965) found no nodules on Ginkgo biloba and, although they recorded Hiltner (1903) as having reported their occurrence, no evidence of this could be found in his paper. Similarly, it was not found that Yeates (1924) had recorded nodules on Araucaria cunninghami. Uemura (1964) appears to be the only person to have reported nodules on Sciadopitys verticillata, stating that they closely resembled those of Podocarpus macrophyllus but were smaller and appeared as ‘“‘ narrow ellipsoides”’. He gave no detail of their structure but reported that Noelle (1910) and Laing (1923) had found them to be mycorrhizal. These authors, however, recorded the presence of endotrophic mycorrhizas but made no mention of nodules. Daugherty (1963) saw beaded roots on the fossil Araucariorhiza joae and suggested that the bulb-like expansions at the tips might possibly be incipient nodules. Such an occurrence, however, has not been observed in living conifers and it seems probable that he was observing beaded roots at the commencement of a new growth cycle. It has now been shown that the nodules of Podocarpus spp. develop as normal features of the roots, are not dependent on any microorganisms for their develop- ment, and differ markedly from short roots, being fully differentiated structures with no root cap or apical meristem and with an endodermis surrounding and overarching the vascular strand (Khan, 1967). Hence, in view of the uncertainty concerning the nature of many of the structures described as exostoses, mamelons, tubercles, and nodules for other members of the Coniferales and for members of the Ginkgoales and Taxales, the survey reported below was carried out. Plants were obtained from a variety of sources in New South Wales, their root systems examined for the presence of nodule-like structures, and sections made to observe their anatomy. RESULTS AND DISCUSSION The results of this survey are set out in Table 2. All species of the Araucariaceae, Podocarpaceae and Sciadopityaceae examined bore nodules analogous to those described for Podocarpus spp., bore short roots as well, produced beaded roots as a result of metacutization and subsequent regrowth, and contained a vesicular-arbuscular endophyte in the cortices of both the nodules and young roots. TABLE 2 A Record of Nodules, Mycorrhizas and Beaded Rootlets Observed on Species of the Ginkgoales, Taxales and Coniferales (+ =present, ——=absent, V=vesicular-arbuscular, E=ectotrophic) Order, Family, Genus, Species Nodules Beaded Mycorrhiza Rootlets GINKGOALES GINKGOACEAE Gingko biloba L. as is ae ie om tL +V TAXALES TAXACEAE Taxus baccata L. 42 Me pis Si3 = + +V CONIFERALES ARAUCARIACEAE Agathis australis Salisbury .. Ah: of ae =| Stal A. dammara (Lambert) L. C. Richard He) a os +V A. moorei (Lindley) Masters bie i a = +V A. robusta (C. Moore) F. M. Bailey es = _ aE A. vitiensis (Seeman) Drake. . ay a + +V Araucaria araucana (Molina) K. Koch ao A. cunninghamii Aiton ex D. Don.. a + A. heterophylla (Salisbury) Franco .. a. at + +V A. columnaris (Forster f.) Hooker + — PROCEEDINGS OF THE LINNEAN Society or NEw SourH Waters, Vou. 97, Part 1 38 ROOT NODULES IN GINKGOALES, TAXALES, CONIFERALES TABLE 2 A Record of Nodules, Mycorrhizas and Beaded Rootlets Observed on Species of the Ginkgoales, Taxales and Coniferales (+ =present, — absent, V—vesicular-arbuscular, HK —ectotrophic) Order, Family, Genus, Species Nodules Beaded Mvcorrhiza Rootlets CUPRESSACEAE Austrocedrus chilensis (D. Don) Florin et Boutelje .. en oe a — — — Callitris eee (Parl.) F. Mueller. . et “= _ WV C. columellaris F. Mueller .. + = +V C. rhomboidea R.Br. ex A. et L. C. Richard — -- +V Chamaecyparis obtusa (Sieb. et Zuec.) Endl. — + +V Cupressus arizonica Greene .. A La — + +V C. funebris Endl. ls Bee ey _ + WV C. glabra Sudworth .. sie ne Re — +- +V C. sempervirens L. .. ag Dy a — + a= \V C. torulosa D. Don .. — + LV Pokienia hodginstt (Dunn) Henry et Thomas — + +tV Juniperus communis L. Hi = == Vi Libocedrus plumosa (D. Don) Sargent ee. — + +V Platycladus orientalis (L.) Franco .. ‘es -— — +V Thujopsis dolabrata (L.f.) Sieb. et Zucc. .. — -- = Tetraclinus articulata (Vahl) Masters id = = +V Widdringtonia whytei Rendle ae ot — — sey PINACEAE Abies nordmanniana (Steven) Spach ae — + +E Cedrus deodara Loudon : nye Be — — +H Keteleerta davidiana (Bertrand) Beissner .. — + +E Larix kaempferi (Lambert) Carriere ae — + +H Picea abies (L.) Karsten os he oa _ + +E Pinus radiata D. Don im ae = ot +E P. wallichiana A. B. Jackson Ae a — a +E Pseudotsuga menziesii (Mirbel) Franco ce - - +H Tsuga canadensis (1..) Carriere oe >. — — +E PODOCARPACEAE Dacrydium franklinii Hooker f. : ae + —V Miucrostrobos fitzgeraldu (¥. apis Garden et Johnson ells ue == \Vf Phyllocladus hypophyllus Hooker f.. at +V P. trichomanoides D. Don .. ai Nes ae + +V Podocarpus brassii Pilger... “ch i ae os ==" P. compactus Wasscher Se a ee a = WV P. elatus R.Br. ex Endl. ie oF oe =f Sig ae P. falcatus (Thunberg) R.Br. alle oe == ale ee P. ladet F. M. Bailey ua i a +5 athe P. latifolius (Thunberg) R. Be, Bi ee at ae Vi P. lawrencet Hooker f. se - + +V P. macrophyllus (Thunberg) D: Don iss _ — =e P. spinulosus (Sm.) R.Br. ex Mirbel ye = + +V TAXODIACEAE Cryptomeria japonica (L.f.) D. Don 5.8 — oS —V Cunninghamia lanceolata (Lambert) Hooker = aL WV Glyptostrobus pensilis (Staunton ex D. Dow) K. Koch = = +V Metasequoia glyptostroboides Hu et Cheng . — +V Sequoia sempervirens (Lamb.) Endl. oe — _ + Vi Sequotadendron giganteum (Lindl.) Bucholz . . = _ SEW Taxodium distichum (L.) l.. C. Richard oe — — = Vi T. mucronatum Tenore = = +V SCIADOPITYACEAE Sciadopitys verticillata (Thunberg) Sieb. et Zuce. ae ue We a at + = se PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, Vou. 97, Part 1 A. G. KHAN AND P. G. VALDER 39 All members of the Podocarpaceae examined bore numerous nodules, of more or less uniform size within a species and usually in two opposite rows, giving the roots a most distinctive appearance (Pl. I, Fig. a). The size of the nodules varied according to the species, the smallest (0-3-0-5 mm. diameter) being those of Microstrobus fitzgeraldu, those of Phyllocladus hypophyllus, P. trichomanotides and Dacrydium franklin being medium sized (0-5-0-9 mm.), and those of Podocarpus spp. being the largest (0-8-1:5mm.). As observed by Sprat (1912) and Baylis et al. (1963), the vascular strand in species with small nodules is very rudimentary, but in Podocarpus spp. it is sufficiently developed to show a diarch structure. Unfortunately, material of Microcachrys and Acmopyle was not obtaimed, as it would be particularly interesting to confirm the occurrence of nodules in these genera. The only record for Acmopyle is that for Acmopyle panchert by Sahni (1920). He reported that the roots of this plant bore tubercles but gave no details of their structure, and his only illustration was of a longi- tudinal section of what he claimed to be a tubercle regenerating and becoming a root. This illustration shows no evidence of the structure peculiar to the nodules of other genera of the Podocarpaceae. In the Araucariaceae the roots of all species of Agathis and Araucaria examined bore nodules analogous to those occurring in Podocarpus, although they were more elongated and arranged in a much less regular fashion (PI. 1, Fig. b). The root systems of these plants, therefore, presented a much less characteristic appearance than those of members of the Podocarpaceae. Only in Araucaria araucana was anything approaching the regular arrangement occurring in Podocarpus seen. However, although the nodules in Agathis and Araucaria were elongated and less regularly arranged, they were structurally analogous to those of Podocarpus, being fully differentiated, lacking a root cap and apical meristem and having a vascular strand completely overarched by the endodermis (Pl. 1, Fig. ¢). Like the nodules of members of the Podocarpaceae, they exhibited regeneration from cells of the pericycle, and it seems reasonable to suppose that they occur throughout the Araucariaceae. The only other plant on which such nodules were found was Sciadopitys verticillata (Pl. 1, Fig. d), the single representative of the family Sciadopityaceae. Here again the nodules were elongated, variable in size, and arranged irregularly. These nodules, too, exhibited regeneration from the pericycle. All members of the Pinaceae examined were involved in mycorrhizal associations of the ectotrophic type and had short roots, each with an apical meristem, open-ended endodermis, and root cap. Several of them had developed beaded rootlets as well. In Ginkgo biloba, Taxus baccata, and all members of the Cupressaceae and Taxodiaceae examined, the long roots bore short lateral roots only and, with the exception of Austrocedrus chilensis and Thujopsis dolabrata, contained vesicular- arbuscular endophytes. Many of them were exhibiting beaded rootlets as well, and it may well be that these occur also in those species for which they were not recorded in this survey, just as A. chilensis and T. dolabrata would be expected to be mycorrhizal under other circumstances. The root systems of many of these plants, with their short roots and beaded rootlets, bore a striking resemblance to those of members of the Araucariaceae. Thus it is not surprising that authors should have reported the presence ot nodules on their root systems and even, as was done by Janse (1897), on angio- sperms such as Acer spp., which have root systems very similar in appearance. As a result of the present survey, then, a clear pattern has emerged with regard to the occurrence of nodules and mycorrhizas amongst the Ginkgoales, Taxales and Coniferales. Nodules of the Podocarpus-type have been found only in the Araucariaceae, Podocarpaceae and Sciadopityaceae, and vesicular- arbuscular mycorrhizas seem to be of general occurrence in all families except the Pinaceae, the members of which form the ectotrophic type. It must be PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Watzs, Vou. 97, Part | 40 ROOT NODULES IN GINKGOALES, TAXALES, CONIFERALES emphasized, however, that most of the plants examined were growing in cultiva- tion far from their natural habitats and only a selection of species was examined. Thus there may well be exceptions to the generalization stated above. As far aS mycorrhizas are concerned, it may yet be shown that some species can form more than one type. Such an occurrence, for instance, is reported by Filer (1969), who records, amongst other observations, that Quercus phellos and Populus deltoides have mostly ectotrophic, frequently ectendotrophic, and sometmies endotrophic mycorrhizas. The taxonomic significance of the occurrence of nodules has been mentioned by Sprat (1912), who discusses the affinities of the Podocarpaceae and Araueari- aceae, noting that they have much the same geographical distribution. Both she and Saxton (19305) regard the universal occurrence of nodules to be a factor lending weight to the grouping of genera in the Podocarpaceae. The occurrence of nodules in Sciadopitys is also a factor strengthening the separation of this monotypic genus from the Taxodiaceae and its placement in a family of its own. This Japanese plant occurs within the distribution of Podocarpus and, according to Dallimore and Jackson (1966), Greguss places it in the Podocarpaceae on the basis of the similarities in wood structure. In the present study it was also observed that when the short roots became dormant the tips underwent the same type of metacutization as that observed in Podocarpus. It is unfortunate that the word ‘‘ nodule’ should have been applied at all to structures produced laterally on the root systems of conifers and their relatives, since it suggests analogies with the nodules of legumes and plants such as Casuarina spp. Had they been given a different name to begin with, it is doubtful whether confusion would ever have arisen in the literature concerning their function and mode of origin. Much work still remains, however, before it can be decided whether or not their function differs in any way from that of other roots with vesicular-arbuseular endophytes. ACKNOWLEDGEMENTS The writers particularly wish to thank Mr. J. Fairburn for taking the photographs, members of the National Herbarium of N.S.W. for checking the names of all the plants examined, and Dr. I. V. Newman for his advice and assistance. References Auten, EH. K., and ALLEN, O. N., 1965.—Nonlegummous plant symbiosis. Proc. 25th a. Bool. Collog., Ore. St. Coll., Gilmour, C. M., and Allen, O. N., eds. Oregon State University Press, Corvallis, pp. 77-106. Bayuiss, G. T. S., McNass, R. F. R., and Morrison, T. M., 1963.—The mycorrhizal nodules of Podocarpus. Trans. Br. mycol. Soc., 46: 378-384. Brcxine, J. H., 1965.—Nitrogen fixation and mycorrhiza in Podocarpus root nodules. Pl. Soil, 23: 213-226. Brevesxi, R. L., 1959.—Factors affecting growth and distribution of the kauri (Agathis australis Salisb.). Aust. J. Bot., 7: 252-294. Bonn, G., 1959.—The incidence and importance of biological fixation of mtrogen. Adumt Sev., Lond., 15: 382-386. , 1967.—Fixation of nitrogen by higher plants other than legumes. A. Rev. Pl. Physzol.., 18: 107-126. BorroMiey, W. B., 1913.—The root nodules of the Podocarpaceae. Rep. Br. Ass. Advumt Scv., 82: 679. Cockayne, L., 1921.—The Vegetation of New Zealand. Leipzig. DatuirmoreE, W., and Jackson, A. B., 1966.—A Handbook of the Coniferae and Ginkgoaceae, 4th ed., revised by S. G. Harrison. Edward Arnold, London. DaueuHerty, L. H., 1963.—Triassic roots from the Petrified Forest National Park. Am. J. Bot., 50: 802-805. Eas, K., and Munpine, H., 1951.—Uber den gehalt an Haminkopen in den Wurzelknollchen von Nicht-Leguminosen. Naturwissenschaften, 38: 548-549. Firer, T. H., Jr., 1969.—Mycorrhizae in twenty southern bottomland hardwoods. Abs. Papers, XI Internat. bot. Congr., p. 59. PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SoutrH Watss, Vou. 97, Part 1 Proc. LINN. Soc. N.S.W., Vol. 97, Part 1 PLATE I i PLATE I (a) Root of Podocarpus spinulosus showing nodules. x 1}. (6) Root of Araucaria cunninghamii showing nodules and beaded rootlets. x 14. (c) Longitudinal section of nodule of A. cunninghamii, showing endodermis enclosing the vascular system and absence of root cap and apical meristem. x 24. (d) Longitudinal section of nodule of Sciadopitys verticillata. 24. ae al nN a) ees ee A. G. KHAN AND P. G. VALDER 41 FurMAN, T. E., 1964.—Mycorrhizal nodules of Podocarpus. Abs. in Bull. ecol. Soc. Am., 45: 151. Hittner, L., 1903.—Beitrage zur Mykorrhizafrage. I. Uber die biologische und physiologische Bedeutung den endotropen Mykorrhiza. Naturw. Z. Forst-u. -Landw., 1: 9-25. Hooxer, J. D., 1854.—On some remarkable spherical exostoses developed on the roots of various species of Coniferae. Proc. Linn. Soc. Lond., 2: 335-336. JANSE, J. M., 1897.—Les endophytes radicaux de quelques plantes javanaises. Annls Jard. bot. Buitenz., 14: 53-201. Kuan, A. G., 1967.—Podocarpus root nodules in sterile culture. Nature, Lond., 215: 1170. McLuckte, J., 1923.—Studies in symbiosis. III. Contribution to the morphology and physiology of the root nodules of Podocarpus spinulosa and P. elata. Proc. Linn. Soc. N.S.W., 48: 82-93. Morrison, T. M., and Eneutsn, D. A., 1967.—The significance of mycorrhizal nodules of Agathis australis. New Phytol., 66: 245-250. Nosse, F., and Hittner, L., 1899.—Die endotrophe Mykorrhiza von Podocarpus und ihre physiologische Bedeutung. Landwn VersStnen, 51: 241-245. Norte, W., 1910.—Studien zur vergleichenden Anatomie und Morphologie der Koniferenwurzeln mit Rucksichtauf die Systematik. Bot. Ztg, 68: 169-266. PARKER, R. N., 1932.—Casuarina root nodules. Indian Forester, 58: 362-364. Perri, L., 1903.—Ricerche sul significato morfologico e fisiologico dei prosporidi (sporangioli di Janse) nelle micorize endotrophiche. Nuovo G. bot. ital., 10: 541—562. Putiiuires, J., 1932.—Root nodules of Podocarpus. Hcology, 13: 189-195. Saunt, B., 1920.—On the structure and affinities of Acmopyle pancheri Pilger. Phil. Trans. R. Soc., 210: 253-310. Saxton, W. T., 1930a.—The root nodules of Podocarpaceae. S. Afr. J. Sci., 27: 323-325. , 19306.—Notes on conifers. VII. Pherosphaera hookertana Archer. Ann. Bot., N.S., 44: 957-963. ScHAEDE, R., 1943.—Die Symbiose in den Wurzelknollechen der Podocarpeen. Planta, 33 : 703-720. SarpaTa. K., 1902.—Cytologische studien uber die endotrophen Mykorrhizen. Jb. wiss. Bot.. 37: 643-684. Sprat. E. R., 1912.—The formation and physiological significance of root nodules in the Podo- carpineae. Ann. Bot., N.S., 26: 801-814. Uemura, S.. 1964.—Isolation and properties of microorganisms from root nodules of non- leguminous plants. Bull. Govt Forest Hup. Stn Meguro, 167: 59-91. Yeates. J. S., 1924.—The root nodules of New Zealand pines. N.Z. Jl. Sci. Technol., 7: 121-124. PROCEEDINGS OF THE LINNEAN Socrety oF New SourH WaAtLsEs, Vou. 97, Part 1 TYPE SPECIMENS IN THE MACLEAY MUSEUM, UNIVERSITY OF SYDNEY VI. MOLLUSCS W. F. PONDER The Australian Museum, Sydney AND P. J. STANBURY The Macleay Museum, University of Sydney [Accepted for publication 15th September 1971] Synopsis The molluscan type specimens, comprising 112 species, which belong to the Macleay Museum are listed in systematic order. These types are now housed on permanent loan in the Australian Museum, Sydney. INTRODUCTION The Macleay Museum at the University of Sydney contains a comprehensive zoological collection including a number of type specimens. Lists of types of insects (Hahn, 1962), fish (Stanbury, 1968), reptiles (Goldman, Hill and Stanbury, 1969), birds (Stanbury, 1969a), mammals (Stanbury, 1969b) and decapod crustaceans (Griffin and Stanbury, 1970) have been published. This paper lists the types of 112 mollusc species. Most of these types were collected on Sir William Macleay’s expedition to the coast of New Guinea in the Chevert (for details of this voyage see Macmillan, 1956). Several molluscan types that should be located in the Macleay Museum were not found during the present search for type specimens and it seems likely that they are permanently lost. Only those specimens actually found are included in the list. Specimens have been taken to be types if the specimen agrees precisely with the original description and the locality is one of those originally stated. The type specimens appearing in the following list are now on permanent loan to the Australian Museum, Sydney. The Macleay Museum also contains the remainder of the Chevert gastropod material on which Brazier reported. The bivalves have not been worked on, and are still housed in the Macleay Museum. The gastropods are of special value because for a number of the species they are the only specimens which have been recorded from Australia. All the specimens in the Macleay Museum were stored dry, in cardboard trays, with or without transparent glass lids. In almost all instances the labels are handwritten and the handwriting is that of Masters, the first Curator of the Macleay Museum. None of the specimens has been previously registered. THE Typr LIst The list of types is arranged in systematic order. Modern generic locations are indicated where these have been changed since the original description. Most of the species were described by Brazier (1872, 1875, 1876, 1877, 1878, 1879, 1881), who did not accompany his descriptions with illustrations. Conse- quently, an indication is given, where possible, of the source of a figure of each species. Hedley (1891, 1901) figured some of Brazier’s type specimens. In PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WaAtss, Vou. 97, Part 1 W. F. PONDER AND P. J. STANBURY 43 some cases it has been possible to identify positively the figured specimen, and this has been selected as a lectotype. This action is taken because Hedley’s excellent figures are generally used for Brazier’s species, so that giving the specimens on which they are based lectotype status will tie them permanently to Brazier’s names. Registered numbers prefixed by ‘‘ A” refer to Macleay Museum numbers. Those prefixed by ‘“‘ C ” refer to Australian Museum numbers. ACKNOWLEDGEMENTS We are grateful to Mr. P. H. Colman, and Mrs. K. Cole and Miss Jeannine Kemple for their technica] assistance in compiling this list. Dr. A. W. B. Powell of Auckland, New Zealand, checked the generic allocations of the species of the family Turridae. PROCEEDINGS oF THE LINNEAN Soorsry or New SoutuH WALES, VoL. 97, Part 1 THE MACLEAY MUSEUM TYPE SPECIMENS IN 44 puidndoanoas (sodAquds) ¢ Spuyys] (AOTZRLE] ISDDUD —= YU srstpoisino) Vyjauidn gy se) #-e Soy ‘er Id “Foe + eset ‘yg p.BIy S9L V OpBIsINO’T “puB[ST Jessoy—=_, vouINy) MON ,, “G FGL8] ‘lover wpbun nurdng 1Aqud 6I-8T s8sy psodvja FT (sodAquAs) cE “On Nel BARE 8 eee ‘tajouny)-ouoivddey, p, Sty PPV ‘IS V poulny) MeN ‘puRls— ern x / ‘TIL 2 PQL81 ‘xaizerge rassouo npaurdne puadndaqu py Ck a = es i Lel V (sodAqguds) 9] PLEL ‘ossoay wpunyod nurdng jo ‘udAg PI) “MOAI MOAROpUT ‘'G + PELE{T ‘letzerg tinaj~onw nurdngy ic nuadndaubiy SEL V [GRl ‘wptaqd sinbuyg vuidng jo udy (sodAquds) ez RIRIISNY “YN ‘purs~ Motregq ‘LE “sy ‘g ‘1d ‘oz : 92061 ‘AeTpoH “q ‘vg s3y ‘oT Id ‘OOT : 898T ‘XoD psy ‘OSL + POLST “dolzerq vpayu nudng (nuednding) nurdndaubrs (sedAquds) ¢ 'g8 “sy ‘e 1d ‘0% : az06T ‘AoIPOH p.31h 9€1 V BYBYSNY “WN ‘pues, wyed ‘ITI-OLL : POLST ‘to1zeag rassou0 nuadng pruovosany dg (sodAquds) Zz 3 a = 9F V “ZO V Rous) MON ‘puR[s— o[n x ‘TILT (= POLST ‘tolzerg wlaxoo piuwoyazy (sodAquds) ¢ FOL V osejediyary Ssuoulojog ‘purysy Aimsrory, ‘p : 9CL8T ‘aoizeag anrydos (ppjajnYo0u,,) DULdvA TY puodowiay (sodAquds) QF SOL V BYyB4SNY “H'N “€ “ON ‘Spur[sy— preureg GIL + POLST ‘tolzerg, wnajonu Durr FT piuovouanyd (edAqojoy) T LEV BouINy) MON ‘MOIR GIL # POLST ‘lolzerg ouwmu purorazy (sodAquds) ¢ eG 921 V RouInY) MON “IOATY MOQRIT 2 ALLET ‘lolizerg nypsowmupu (wovagdrsa Nt) DUIYALA NT 8h V (sodAquds +) ‘861 + PLO6GI “AotpoH “jou 9ST ‘8IqQteAnog HyD]JAjs DyjaMWolY Fo ‘uAg BI[BIISNY FO ISBOO "WN “€ “ON ‘SpuvysT preurwg ‘LE St YLLST ‘tolzergq pnypUsLo HpJagnwuozs (sedAquds) 1 LQG WV WBIG Semtoy, “s~T Aopureq (sodAquds) 9 OGL V PVI4Y somo, ‘fooyy Iowre A (edAquds) J TOQOeCL V woulnt) MONT ‘MOIR (sodAquds) 7 "8h Sy ‘19 1d “LIF + 688T ‘Arqstid psy OST V elelisny “N ‘yaoX odep ‘PP i CLLST ‘aetzerg wypjnonw ny0)nY,T, (sodAquds) Z ‘of “Sy ‘LT ‘1d ‘821 + PLO6T ‘AoTPOH p.siq GLW BY[eIysnVY JO 4SBOo “AN ‘€ “ON “ST paeureg "ep i CLL ‘aotzerg snsounib snynounzp (sedAquds) 1 O9T V WwVIJG seqoy, ‘pussy Aopureq ‘9% “SU ‘EOL : PZOGT ‘AOTPOHL 7212706 vppeungoung S0J0N 10YIO ‘uAuoudg uUMOUY ‘pedueyy Jl euleN ollouer) ULepoy ‘I9qUINN Jo4stsoyy nv ror UOlZeUsISe(T ‘sueumtaedg jo r9squinn ‘Aqryeoo'T aoUSTEJOYy ‘peqiioseq] YoIyA Tepun oureN PROCEEDINGS OF THE LINNEAN SocireTy or New SoutH WaAtEs, Vou. 97, Part 1 Ker) RY R AND P. J. STANBU , u F. POND W. (odAguds) ] WRG Ssonlop, “puvslsp~ Jo (sodAquds) ()T JB4Q SaLloy, ‘pues, Aepurecy “(ELSI “U0 NF ununuaun “9 uou) “LIE DfLL&T ‘dower, wun wniynday o1d *Aou W0U $e SY ‘OSPF + 6681 ‘AoTpoH wnuuissyunbaja wneynse— UWNLYILMAIOLD) (edAjoqo0[eaed pus edAq0q00]) Zz *(odA49404007 Se uesoya) Fe -3y ‘LT 1d ‘9ZT = PLOGT ‘AOTPOH pS IL V eoulnyy) MON ‘MOIR YT ‘OTE i SLL8T ‘dolzeag, wngniaaiqqn wnrypsaZ -(edA40}00] Sw Uosoys) (sedAyoqooyered puev odAqo0400]) g cz sy ‘LT 1d ‘LET : PLOGT ‘AOTPOHT P.S1 GL V WRG somos, ‘puvsy Aopureq: ‘LIE = fLLST ‘aotzergq wnusafypunor.g wnaysag (sodAquds) ¢z WIPL V voulns) MON “MOZBY (sodAquds) F 5) i 2 ByBYSNY “HN “yIox odep “LI ‘OT SSY ‘G68 + OG6T ‘UOIeSe'T p,SIT (sodAjuds) "LET + 1061 ‘AOTpOH “Jou Iftl V BYBysny YON ‘epiAuery odey "80 : ALLS ‘tolzerg, sapurpins nurossay (odAquds) | CHL V BIBIISNW “N ‘olf Auary edey jo ‘spurysy ow py (sodAquds) QO] ET “SU ‘E6E + OGG ‘UOLOSeTT p.diy ICP V ‘LOG : ALLEY ‘aolzeag, wpjayojnd nurossry BYBLISNY “AN ‘€ ‘ON ‘puvysy preuieg (DIDIDUhZ) nUurossry PrL V (sedAjoqo0;eaed pur odAy4o04007) 9 WRG somoy, ‘puvysyT Aopureq: “(ed 4409007 sev UosoTa) 8c “SY ‘LT 1d ‘221 * PLO6I ‘ACIPOH P.sl "LOE : BLLQT ‘aolzergq suuwaur nurossr1y (n)JavYyILa0 fT) DUL0ssray Ghl V (sodAgoj00;ervd pue odAyoq00/) © BlTBlgsnVY “AN ‘ “ON ‘SpuRs[T paeureg *(ed440}90] Se wosoyo) 96 SY “LT 1d ‘ne : PLO6T ‘ACTPOH P.8A ‘99€ : bLL81 ‘aotzeagq, vqwoiIffo Dvur0ossray Suisstut suouttoods pur[s— ssouesuncy DYAVZIMONYIS SPL V (sedAquds) ¢ JlIBIZQ Soeddoy, “puB[sST ong 63 “SU ‘LT 1d ‘8ZL : PLO6I ‘AIPEH PS "89g : ALLE] ‘aolzeag, Mnowdsuoour purossray (DJJaULsOY T) DULOSSIAT (edAqo400Teaed) | NL OPT V WBAYY Solo], ‘Joo, IOLITe AA (sodAyjoqo0[e1ed) z OPT V PIO WOR odey ‘Avg pny (sodAyoqooyered puke odA404007) ¢ TORI V woulny) MONT ‘MO9BY *(odA40400] SB UoSoYyd) (sodAqoqoopered) z Le ‘Sy ‘LT 1d ‘LeT : M1061 ‘AoIpoH Psu 9FI V JlBIZG Selloy, ‘pus, 9G ‘LOE : GELS ‘atolzeargq savaz purosseay 69 V (sadAquds) 9 G98T “UOAIT, s2xQV.G8nNM HIqQQDX) Jo ‘UxG “ACO'N ‘30019 Ulegseny 6 2 0GLET ‘adolzeagy vuaphy nioyng n0d070 NT (sodAquds) ¢ POL V PIO ‘tary vuIgueurelcy ‘13S: BLET ‘lotzerg, wwosen nundiar 4 “LI-GT s8y “FE Td ‘gon + MZ06T ‘AOTPOH PSM “SIT +: POLST ‘roizeig wnajnu snaoydojohy oad *Aou “uIOU puUasibouuy (sodAquds) 40% "2ET L&E V wipRngsny oN ‘yaoR eden PULLEY ‘lolzeag vawoppaq (sadoumaq) snsoydopah) PROCEEDINGS OF THE LINNEAN Society or NEw SourH WatLEs, Vou. 97, Part 1 UMW (edAyofoy) 1 ‘py SY ‘9T 1d “E21: PLOGI “Ae|poH pi GZt ¥ ywaajyy soatoy, ‘pussy Aopuaecy GLI t PLL8L ‘dolzerg psnfuoo xaunjpy PEL V (edAzojoy) 1 FEET ‘Aqdomog sruysoujoas vaimpy Jo “Usg qveryg setioy, ‘pussy Aopurecy ‘OLT + PLL8T ‘dolzeig, sniwira wainpy Suisstut suouttoeds puvs— Aotod DAP N (sedAquds) ¢ G8 W PIO “0A odeyy (edAguds) { GOL V JLBIQ SoLtoy, “opis “MM ‘Joo yy TOMI AQ (sedAquds) ¢z ALZ& V ‘G8 V ewoulny) MON ‘MOIR YT Il ZS W (edAquds) | "8 “3Y ‘9T Id ‘FZ1 : PLOGI ‘AoTpEH p.3iyq CERT ‘Sopond vurnoajow *O Jo ‘uAY ytBI4g somos, ‘pussy Aopurecy ‘O&S + OLL8I ‘s0lzerg, Dydisosur vyaqunjop iC ‘OL “89 ‘9T 1d “F2r + PLO6T ‘AoTpoH p31 3 auaihg (sedAqudAs) + (OF “CES-1ES S V8 Ve SS SECU eegien Aguas Sonim “eeyerenrg pnopnd (nphwy) nyequnjop E Suisstur sueuttoods pur[s— ong = SuIsstun ri SB S][eys esoyy peqoder (EZT : PIO6T) Aog[poH at | scl V (sodAjuas) &@ “PLI = (FPST ‘spulp{) snapnbyup uopaparn Jo “udg jeg solo, ‘pues, Aopureq :PLL8T “dolzerg aspbun (snwoipidy) wnruopay, < = == = — — = uoiydiiosep [BUIsII0 34 Ul UEALS SUOISUBUIP a4} YIM sede you Seo([ : ‘(4eT -d) 061 “3y S| snsnfuny _(odAquas) I ‘8CT + G96 ‘loluqey pue uosiydoeyy p,31yq = 66) Vv. Blpeagsny yynog ‘Aeg ueyoms) ‘9 + 9GL8T ‘solzerg wpoUpaq snwoupridy Zz es ee ig sanUUyod a (QAveY Dsoporvap DIYDN' JO plooot (GEZ + OLLST) 8,dotzearg =) (sodAqewed) z ‘9 “By : 6S. V {BQ souoy, ‘pussy Aopurecy ‘6G Id ‘9E1 + F261 ‘Ao[poHH wnayjzueshqn, 429) a) 2 ee eee BI (edAyojoy) | D OOT WV JBI Somtoy, ‘pues, Aojureq "G8G : ALLS ‘lolzeig sepqnwn nun a (edAqojoy) | 1% “3y ‘LT 1d ‘GZ : LOGIT ‘AolpoH, p,31yq Ps 10L V WAG souop, ‘pussy Aepusrec, “C8G + ALLE “lolze1g suapu Dwyng : sajsiddvT (sodAqoyjoo,ered) ¢ (1ST V Bounty) MON ‘M09BY ‘(ed.440300] se uesoy—) LSI WV ed A40309]) | ZZ SY ‘LT 1d ‘931 +: PTOI ‘AoTpeH p.31q LIS, ‘udMILCE wisupupdas oqang, Jo ‘uAY 1BIZQ SetLoy, “‘puRls—, Ioq G1E i SLLET ‘solzerg vyDU2Wv9014 sido1joYyors 7, (edAqoqoy) | €e “3Y “LT ‘Id ‘9ZI = PLOGI ‘AeTpoH p,31q OCI V JIBIZG Selloy, ‘pussy AopUrecy STE 3 JLLET ‘olzerg Dzuaprons6 8UAOMOYIULT, © = = Soe eee = é = SeJON IOYIO ‘utAuoucg uUMOUY UOlyeUsIse(y QoUdLEFOYy ‘podury,). Jl euleN oLilevex) UWlopoyy ‘suouttoedg Jo 1aqumy ‘peqiioseq] yoy aopun ourey ‘raquinN 1o4sisoyy ‘Aq B00] TH WALES, Vou. 97, Part 1 y PROCEEDINGS OF THE LINNEAN Society OF NEw Sot bt W. F. PONDER AND P. J. STANBURY T6rL V 6FL V (edAqoqoo[ea1ed) | RByBysny “N .“yaox odey (od 4904901) | Weayg seatoy, ‘pue[sy Aepurecy 0% °“3y "192 (edA440400] SB uosoyo) ‘ol 1d ‘eZT : PLOGI ‘AOTPOH P.Si > PLLST ‘aoizerq wuyoynd njoushcy pyauydnpopnas q (edAyojoy) T "gst ‘Sy ‘9¢ Id ‘oce : zz6T ‘AoTpoHT p.sta SLWV eoulny) MON ‘MOJRST "LGT = POLST ‘atolzerg ~hnsupns vyaunywID PRST “SpUulHy S272Qap 07]24NYIDID se (GGT = POLST) telwerq Aq poyiquep] pyjauydog (sodAqguivd pue edAyofoy) F ‘091 “3Y OLIV qledyg somos, ‘pussy Aojused ‘eq ‘1d ‘L7G 32261 ‘Ao[peH wnuysao nyauydog nyjauydopopnas q (sodAquds) F 891 V yer Sotto], “pussy Wed (edAquds) | "SLIT “Sy ‘eg ‘Id ‘epg = 261 ‘AOIPOH Psa GEL V BW somos, ‘@ ‘ON ‘SpuR[sy prvureg "1GT = POLST ‘lolzerg, wpunudmg DyainywIp ‘Fel “Sy ‘TG ‘1d ‘OTs + ce61 ‘AoTpoH puopydnysopnas q (sedAjuds) [e ‘eo, “3y ‘T ‘Td ‘9gz : F8gT ‘uoAIT, psy OL WV WeRyg somos, ‘pussy Aopureqd ‘PEL 3 POLST ‘aotzearg, ehayusmp vpjaunyInIO (1alze1g) 1070904) njowoo0sbut nypanuraT (sodAquds) ¢ ‘e “Sy ‘or 1d ‘cer : P1061 ‘AoTpOH PS 6L V PIO ‘puvpsT Wied "SET = POLST “tolzerg 10700042 17794NYIMI) (691 * POLST ‘tolzerg Aq ‘GPRT ‘sA90Y DIDULLDIWUNIG DyeUydoG se peytyuepT) (edAqojoy) 1 ‘TOL ‘39 991 V qyeayg Settoy, ‘puepsy Aojureq ‘ge “jd ‘Lzg + 2z6T “AeTpeH wloacyo vyouydog nappa (sodAquds) ¢T TA 691 V youlny) MON ‘MOBS (sodAquds) FT A691 V qIeIAg sedoy, ‘puetsyT Aopureqy (sodAquds) ¢ MT 69T V elperqgsny “N ‘opftAuerg ode (sodAquds) Z ALG9T VW alyeaysny “AN ‘Avg o9j0[1eyO ssooulrd (sodAquds) Z 1 69T W Weg solo], “pur[s— Ia (sodAquds) ¢ ‘By ‘or 1d ‘cer : PLO6T ‘Ao[poH P.StA T69I V eypeqsny “GN “0X odep “LOT + POLST ‘tolzerg hmejonw nyainyIn7O opesanbuy L6V ; ; 888T “YqTUIg (sodAquds) % "sg “Sy ‘g 1d ‘z8p + 8061 ‘AOIPOH P.SMA sisuauojburssa (niypiiqr) nuojoimarg jo “ukg WRIYG Soloy, ‘puBS]T TOI AA "ECT 2 POLST ‘dolzerg wuwagspw 02770LG Lonsinbuy (*‘SOTIITBOOT (sedAguds) 9 Jo 4ST] [BUISII0 ey UI poyou you uoeUTtoods 186 V qieiyg somoy, ‘pussy Aopureq Stuy, “pemsy pur[s—~ Suoy wrory uourttoodg) (edAquds) | ‘1 ‘By ‘or 1d ‘cet + P1061 ‘AoIPOH Patt 86 W qeIq4g soto], ‘pussy 94 "POT-EST + POLST ‘ToIzerq wuepyods Hyp4q *(9d 4409907 Se Uosoyo) Lopsinbuy (sedAjoq00;e1ed pur edAyoqQ007) ¢ IF “99 Gel V aijeagsny uw1eqso\ ‘Avg syareys ‘ep “1d ‘0G 2 2Z61 ‘AoTpPOH Vilatdwnp nuowpa yy PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WatsEs, Vou. 97, Part 1 THE MACLEAY MUSEUM SPECIMENS IN TYPE 48 (sedAyoqo0[eavd puv edAjzo400]) -|- Qe *(odA}OJOo, SB UoSOYd) 6¢ “Sy ‘21 ‘Id ‘OET =: TO6T ‘AelpeH p, Sip G9 V }IBI40 Soltoy, ‘pussy Aopuseq “GR: WELLS] “lolzeagq siswahapuwmp ship (adAqoqo0y;ered) 7 ZOT W eledysny “N ‘et[tAuecr) odey ‘(edAjo}09] se Uosoy—) (sedAjoqo007eavd pure odA40409]) ¢ sé sy ‘21 Id ‘ost : 1061 ‘Aetpery p.si 1 ZOT WV ytioK ody ‘asesseq Aueqry ‘FPS-ES = WLLST ‘lolzeag nsooap now zy — (peyeoo] you usuttoeds peansiy) (edAquds) | “el “3 ‘9% “Id ‘OTE : 0061 ‘Ao[poHL p.S1y LELW WRN4Y solo], ‘pursy Aepurecy ‘PS } WLLST ‘dolzerg DyoLYsinu sapunydoogy . ‘(adA4q0}09] se uesoyo) (edA4o}00]e1ed puw odA40900]) Zz Le Sy ‘LT 14 ‘ect : 106T ‘ActpoHy p, 31g OFL V WeIZG saiio0y, ‘pussy Aoyureq ‘6L-SL 3 WLL8T “tolzerg, Mpoorsshiqn nynorburay *(adAq0}00] se uesoyo) (sodAqoqooy;eaed pue odXy4o 001) ¢ er sy ‘ot ‘Id ‘ect : pTO6T ‘AolpeHT p.Sty SéL V VVN4G soiioy, ‘pussy Aopusrec 626 = PLLST “lolzerg nunj]9 vriwo0EjsopeE Suisstur suouttoeds puvys] Aoyurecp LELV (sodAquds) ¢ BIBIISNY “N ‘yao KR ode y “LI “89 “or 1d “ezl : vTt06T ‘Aolpor p.siq ‘68E = PLLST ‘toIzeI1g suaffp nuuozsopcE L€L V LL8L ‘doizergq njydiwoo -GQ Jo -udg (adAqojoy) J WIIG soos, ‘puelsy Aopurecq "81 “Su “41 ‘1d “ezy : vT061 ‘Aoipory psig ‘096 = PLL8I “lolzerq j20d nruuojpsopC O&T V LLST ‘dolzeag, mdwoo *G Jo “Udy (sodAquds) 7 }'aIqQ setoy, ‘pussy Aoyusrecy ‘6T “Sy ‘ort Id ‘eat : -L06T ‘AolpoR p.3t¥q ‘O09G > PLLST *‘TOIZVIg, pynawod DUWOISOP() (sodAjoqoopered pue edAqo4o0]) TT] ‘(odA}0}00] sve uosoy) 91 “8y “ot Id ‘e2t : P1061 ‘AorpoHy p.Siy 6L VW WRG Somos, ‘pus[sy Aepurecy ‘09% + PLL8I ‘totzeag, wydwoo niuuojzsopy: : ‘(od4q0400] se uesoyo) DIPS (sedAyoqoeyered pug edAqoqo0[) ¢ FI SY ‘OT 1d ‘e@T : MTO6T ‘AoTpoH, p.3iy 8 V Bounty) MON ‘MOBY ‘SEZ } PLLRT ‘tolweag, wuydn nyrwoqury, DIZPUULAY CS) (sodAjojoo[ered pus edAy4o0309]) 9 : ‘(edA}0400] SB Uesoyd) Zt 3y ‘ot 1d “beL = PLOT ‘AoTpeH, p.3Iq hh, WV4G Soros, ‘pussy Aojusecy ‘LOZ : PLL8T ‘tolzerg sesualiayusop nppruoqguny, ‘(0981 ‘Stuepy “Vy uou) LOG : PLLET ‘dotzerg nwixa +7 ord -aou ‘urou oyvosbih (edAqofoy) 1 fail ss CV RIBIMSNY “HN ‘Z ‘ON ‘spuvysy Aorog ‘OT 1d “PZT + PLOBL ‘Ao[poH Ywaneyo vppruoquny Ssnasrbih J (edAyofoy) J ‘99T “SY “86% : 6G6I ‘UoroseT p.S1q TW JIBIZQ Somtoy, ‘puelsy Aojureqr ‘6GE-8EE = PLLET ‘selzerg vsnfuoo vyuoqung, SeION LIYIO ‘urduoudg UMOUY, ‘posuryy) Jl eUleN olletes) WiopoyT “TeGUINN .oySisoeyy uolyeUsIsecy ‘sueuttoedg Fo aequiny ‘AQITROOTT aOUD.LOFOYy ‘poqiuoseq? Yor aepun ourerny PROCEEDINGS Or THE LINNEAN SocieTY OF New SourH WALES, Vou. 97, Part 1 W. F. PONDER AND P. J. STANBURY DUISSTUL OLB suottinods purfsp ssouosun¢p puR puURIST ony (opr Masry) DIdCd01SY}) WPEL V UPEL V &l V (sodAquds) ¢ }IBLIY Solo, SpuB[s]T LOLLTR AL (sedAquas) ¢ PIO (N ‘pues, uvodoy (sodAquds) T¢ PIBIFY Sato, “‘puefs] Jog 6€ “S315 “6 1d “GL + GZ06T ‘AoTpEeH psi (npnUeq?DUsrp) YjIdoI01jsD}) (sodAquAs) F ‘OIL «¢ POLST ‘selzeag ahvapnu (obysa,) vdng GElV “MON ‘olrenboeyy, oer] “6I-SI = PEL8T ‘toelzeag dapsso. (obyway,) vdng (sodAquds) QT —_ — TGS V PIO ‘xt0R ede) ‘purjsy Aueqry CGLV (sodAquds) 9] OLST ‘talzeigd iswapspiue YurpjpayoUloy, Jo *UAG Byetysny “WN ‘spueys] eurozy ‘601 = POLST ‘dolzeag dappivuasb vUtp{ajNULO, 7, (sdourpjanusof,) vapiyjauny SST V (sodAquds) FT VB4G SoMoy, ‘pussy Aopureq. “PI-EL “83u ‘PE Id ‘cOL : MZO6T ‘APTPOH P. Sly “SOL 3 POLST ‘lolzerg, Usuaysnul DULIIAIDULO,], SDUUSD) Y ccl V (sedAgudAs) + RouInsy) MON ‘puRys] o[nA “601 ! POLST ‘dolzerg?D stiupsatiay DWL]10IDULO SDUUSD) A ISl V Blpelgsny (sedAqguds) < “ODN “€ (ON ‘pue[s] paeured “er 3y ‘PE 1d ‘cOL 2 OZOGT “AC[POH P.Stu ‘OLL «: POLST ‘dlolzerg stuoyona nUrjjaj0ULO 7, uinesnyy ysiiig ut edAqyojoH DUALDBODI A 191 V BIfeIsny (sedAgeared) g “AN ‘pueysy Aorz91y ‘16-61 ‘$3 “€8 1d “OL9 * 4GL8T ‘IoIzelq wpunymobh nurnuuojdig “1 + PELST 9EC6L “) (sedAqguds) Qe BIROSNY “AWN ‘foalyp paeuuary “€1Z 2 SPELL ‘eyeporyl wueimmmos vauuydag psnjay 88 V ORR ‘UosPE MA srjsoziydiuy snynoiuQ jo “UAY (edAquads) [| JIA SWOT, ‘pueysy Aopurecy (ZL8T ‘Swepy “HH Pinu “9 uoU) 18-08 ‘1elzelgq vinuau puyouhy oad *-Aou “utoU CTE 3 €681T ‘Auqs[id tarznug nuyouh, 2 MOLLST puyouhg (sodAquds) $9 LOW qleigg seioy, ‘puvysy Aepurecy ‘L8-98 : WELLS] ‘loizerg wsroqnp ship puyosyhy (sodAjuds) $F] €ZL V VBI Solo, ‘pusysy Aopurecp ‘TR i WLLST ‘doizerg vsounibh nysaupy puyouhy (sedAquds) ¢z 89 V {B49 Solo, ‘pue[sy Aopurecy ‘Q8 i UWLLQT ‘toizeag, nayognd shjp (ed&yojoy) | 99 V We Seioy, ‘pussy Aepurecy "OR 1 WLLST ‘lolzerg psiap ship (edAquds) { 1f9 V viferysny “N ‘oy[tAuesp ede) (sedAquds) + OL OF “BY ‘LT 1d ‘Oe + PTO6T “AeIpeEH p.S1y1 ie WeRzpQ soawoy, “puepsy Aopurecy aa “OR—GQ MELT RT Sdolzeig, tpaaayo shyp 97, Part 1 Sociery oF New Sourn WALES, VOL. PROCEEDINGS OF THE LINNEAN ee ee FMI IT ne SE EE MT a eT A oe ne Re tbe ho BA FACE I Sea a OE OLMIS tt rE - : ; sane soeeeneesiateneieeseneeienenee:emmmmmmsiciiteiibeneiemmmminimmeaiiiamiiammaiaaiaa ice <4 4 hue PAH pt Soar = si EARN ALLA LTT ITS TOU a aed Temersied 6 “1d “GL =: T681 “Aeqp: mk pst GIL V routny) MON “PURIST OTA “SOL : POLST “ToIZergq dayings (snynuop) xYaFT sushoode iy (sodAquds) ZT F fol \7 nouns) MON ‘puRIs] oT A ‘OOL : POLST ‘dotzerg, oyddns (nissmoyy,) wyazy (sodAquds) TT] ‘FI 3y 106#9 “0 BITeIIsnVy “M'N ‘osuey Jomreg ‘e ‘jd ‘pF 36861 ‘oTepoay snquay sushonuysa jy TYPE SPECIMENS IN THE MACLEAY MUSEUM 50 (LOJUOADA) 4090NT (; odAguds) [ *T68t ‘Ao1peH Pst ‘OOT * POLST ‘toIzerg snynwun (nissHpMYy,T,) xyazT ILI © qeIY® Solos, ‘puR[sT ssoussuncy (sod&quds) 6] ‘OT “3y ‘Or ‘1d ‘8 cP V ‘OL V woulnty) MON “IOATY MOVES Ic wv (;, sod4yodo4 10 sodAqeaed) g (G98T ‘Wossnoy) srsuaowns nyapapvyT jo “ukg SepLIqoxT MONT ‘uInoyIouUW ‘9-7 ‘s3y “ee 1d ‘96 : 6S6I ‘WIEIOg p.suq ‘P8-E8 2 OL8I “X0D vmMJoIUD KAT DAOMSALLO, J, (edAguds) J ALGIT V WIVBI9G SaMoy, “puBIsT TOTIIe AA (sedAquds) OT Mm GTT V Weg Selo], ‘pussy ynu0s0/) (sod&quds) gT "etl W WRG soto], ‘pueys—T ong (sodAquds) £7 TCIL V WeyS Selo], “purjsyT oq (sodAquds) g¢ ‘TI-6 ‘s3y ‘Fe 1d ‘cor : oZ06T ‘ACTPOH PST cll V PIO “y10R odep “EOL : POLST ‘tolzergq wurpyods (nynoq) Tyo snwopilan?) (edAqojoy) T 801 ¥ vI[erysnVy ‘or “Sy ‘eg 1d ‘Tz : 92061 ‘AOTPOH pst ‘AN ‘oyftAuery odey yo ‘spue[s—T owop ‘POL : POLST ‘totzerg 2a7p1aUasb (snynuoy) x19 FT Dysapunjaqy (sodAquds) “6I-LI “s8y “Z Id ‘oT + 9T06T ‘AOTPOH, p.SiT 6IT V PIO ‘yepudey = °F_—-:: OGLE ‘Aolzerg__ sresuaynpuhinb (nesswyoy,[,) wea FT (npbayndwuy) nuwosidng (odAqoToy) T “FOL + POLST ‘toIzerg sod (snjnuep) xyaH *(edA40900T se Uosoyo) It ‘sy ‘or ‘1d ‘6z : 1681 ‘ACTPOH Ps ‘TOT ‘rolzelg, asysaqiop (nprydoyzyoQ) «yor O1L V eIeysny “N “puelsy Aueqry SNYIOLDAD T (edAgered puv odAqo0q00e]) Z 86 V goulnt) MON “pueysy on > POL8T nurdowiu Fy (sodAyuds) Z ro V woulns) MONT *pURIST o[N (Ajoatqoedser 968F9 ‘L68F9 *O) (sodAjered pue odAyojoy) & eledisny “AN ‘osuey szcideN-doieg "€2-2G ‘8 “3g ‘9z ‘Id ‘Lee : F6RT ‘ACTPOH P.Sla "SOL 3: POLST ‘iotzerq whnajonu snwyng *g “Sy ‘tT ‘1d ‘IT :6€61 ‘oTeperyT vIdnuup vdndowey 7 SOJON IOYIO ‘urkuoukg UMOUYT ‘pasueyO jl eureNy ollouex) UEpoy, ‘TaqUINNT IoysTsoyy UOlyVUSISECT ‘sueutioedg Jo requInN “Aqyeoo'T Q0Ua1OJOry “peqiioseqd YoIyM JepuNn ouIBNy PROCEEDINGS OF THE LINNEAN SociETY oF New SoutH Watss, Vot. 97, Part 1 (sodAgvaied pus ad &4oj0u) $G (sedAgeaed) -9e6L *) BITBIISNY “M'N ‘eduey «vos— (edAgojoy) LEst9 ‘e ‘sy ‘p 1d ‘09 : 66L ‘O[BVpoal VNuZsU0D DpHboYyaT (sodAquds) ¢ ; “G-p say ‘OZ ‘Id ‘99¢ : 2681 ‘X0D tquep (vupoy]) xyaH ‘91 “Sy ‘e Id ‘1G = 661 ‘O[epoay D2100U2 nryonuyDWYWSa MY "al ray) ‘¢ ‘1d ‘1G : 6861 ‘e[epery Use DYyoDIID.LYWS9 "SL “8 ‘e 1d ‘gp : 661 ‘OTepod] MpO0WOW HIY90.19089.10,], "66-86 * POL8T ‘lolzeig zewappaq (vpyfiyy) xyoH E88F9 BIBIISNW “MN ‘osuey Joltieg (edAyojoy) [| 988F9 “) BIBIISNY “AA'N ‘osuey rowed 8 (edAyofoy) | 888F9 “OD eBlelysny “AA'N ‘osuey dowieg g aaa ws (sedAqeared) Z €996L ‘OD BypBaysny “M'N ‘osuey (sordeyy) somreg a pyanydin ay (sedAjuds) OT 9II V PIO “y40A odeH ‘puvsy Auvqry (1g81 (sodAqevaed) + ‘sequo) susuaipunpyund, vyjanydinpy jo “uAdg PIO 'N ‘puvysy wyed "66 ? POLST ‘toIzerg isawnl (opyuhyy) xyaH W. F. PONDER AND P. J. STANBURY pysabuviyy) (4 odAqvated) | ‘PI-ZL S8y ‘L 1d ‘LE : S061 “IeGOM psy €LIL WV Bijeaysny yo, ‘sesuey AqgjoT yunop 819 : ZLST ‘dolzerg 2aynvb (sajuoz) xyazy snphysoov (sodAquds) z ‘Pp 3 0CL8T OL WV osejediyory suoULo[og ‘Ieizeig, wpunpiofopu (snjhzsoovamngy) snuwyng eiygdjopeptyd “Og “yeN “peoy ur edAzojoH snyhqsoov 1g (edAqered) [ 091 VW BIUOpEeTe MON “MA'N ‘Avg ouoyoN "98G = [88 ‘lolzeIg twapsscu snwyng *(Aurozeue) Fr -3y ‘6 ‘Id ‘¢ “By ‘g Id ‘*Z -3y ‘9 “1d ‘(Tfeys) SNI8YPOIt) (sedAquds) ¢% 1-¢ s3y‘z ‘1d ‘16 : E88T ‘ujousp-ouoieddey, p.31q7 0G W ‘98 V BOUINL) MON ‘pUBIST INA "TOL : POLST ‘dolzerg, auwowo7 (snosiq) xyazy Djajppoy (sodAquds) 1 ’ 601 ¥ WAS solo, ‘puws] yNUo0I0;) “Gb-OF SBy ‘g “Td ‘OZ + 92061 ‘ACTPEH P.31q (sedAqudAs) 9 COT T60I V BIG Soutoy, ‘pussy ueodon :MOL8L ‘dolzeigr seswaunedau (snynuop) xiwWazT ‘pr 3y ‘e 1d ‘1S + 92061 ‘AIPOH P31 DyaqDpoy (edAjofoy) | COT 901 V RYBIISNYW “AN ‘€ ‘ON ‘pueys] parvusreg :DOL8I ‘lolzeig siswapupusmg (snynuop) xWAzT ae ‘ep “Sy ‘e 1d ‘1% + 92061 ‘AoTPEH PSI DIa]DPOs! (sedAquds) Z ZI LOL WV HBG soqtoy, ‘pussy Aopureq : DOLE ‘lolzeiq sisuahajzusp (snynuog) xryazy pyayppogy (sedAquds) 1% ‘cp “3y ‘¢ “Id ‘0s : 96061 ‘AoTPOH P34 Ill V War4yg solog, ‘pussy Aoyureqy *ZOI-1OL + POLL ‘serzeag, zopaou (snynuop) wyagy puryebang (sedAquds) ¢T ‘9 “By ‘6 1d ‘ex + 1681 ‘AO[pOHT p.siy4 cy V BOUINY) MON ‘puweysyT 9/1 X “TOL? 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F. PONDER AND P. 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GL i WLL8T ‘totzerg AqsoMog snuiiayond uonyp se peytyuept ATsnotseid 6El V (edAyofoy) T qrei4g souoy, ‘pursy Aopusred “OF “LZ ‘Gs S8y “6I “Id ‘TST : OZ6T ‘IIMH pus o[epoay suajjaoxva xnjdosshyy 69 V (sedAquds) + pI ‘uostueqd 4410q ‘TI-OT ssy “C8-Z8 : [881 ‘SpoOM-UOSIUOT, DPIpPLo0s HI70119SN UINesn AL “gsny ul (eg919\)) seddqzered pure (79819 “) edA4qo[oH OF V (sodAqeaed) 62 elpedgsny “MIN ‘toAty paxreuue’T ‘g “By ‘9 ‘1d ‘¢ ‘sy ‘¢ Td ‘TL : $E61 ‘olepouy Mabbouf nyjan1oT g-¢ ssy ‘g “[d “er : 8961 ‘rummy, pure yous, Aq uesoyo edA,oq00'7 (m297]2]09094)) 1191)2'T 69 V ee (edAqoq007RA1ed pure edAqoq0e7T) Z 2 mounts) MAN ‘punog [ley ‘2@ °8y ‘TT 1d ‘88 : 1681 ‘Ae[peH Pst “SOT-LOL 2 POLST ‘telzRag owaz (sn1y901400/)) XIAFT Ss OF THE LINNEAN Society oF NEw SoutH WALES, Vou. 97, Part 1 1 PROCEEDING DA TYPE SPECIMENS IN THE MACLEAY MUSEUM References Brazier, J. W., 1872.—Descriptions of six new species of land-shells from Australia, and Lord Howe’s Island. Proc. Zool. Soc. (Lond.), 1872: 617-619. ————, 1875a.—Descriptions of eleven new species of terrestrial and marine shells, from north-east Australia. Trans. Roy. Soc. N.S.W., 1874: 29-33 (issued separately, dated 1874). , 18756.—Descriptions of eleven new species of terrestrial and marine shells from north- east Australia. Proc. Zool. Soc. (Lond.), 1874: 668-672. , 1875c.—Description of fourteen new species of shells from Australia and the Solomon Islands. Proc. Linn. Soc. N.S.W., 1: 1-9. , 1875d.—Descriptions of eight species of Australian and Tasmanian land and fresh water shells. Proc. Linn. Soc. N.S.W., 1: 17-20. , 1876a.—Descriptions of thirty-five new species of land shells from New Guinea, Australia, and islands in Torres Straits, collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 1: 98-113. , 18766.—List of land shells collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 1: 117-133. , 1876c.—Description of a new Pupina collected during the Chevert Expedition. PRcc. Linn. Soc. N.S.W., 1: 136-137. , 1876d.—A list of the Pleurotomidae collected during the Chevert Expedition, with the description of the new species. Proc. Linn. Soc. N.S.W., 1: 151-162. —, 1877a.—List of marine shells, with descriptions of the new species collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 1: 169-181. , 1877b.—Shells collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W.. 1: 199-215. , 1877c.—Shells collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W.. 1: 224-240. , 1877d.—Shells collected during the Chevert Expedition, with descriptions of the new species. Proc. Linn. Soc. N.S.W., 1: 249-261. , 1877e.—Continuation of the Mollusca of the Chevert Expedition, with new species. Proc. Linn. Soc. N.S.W., 1: 283-301. , 1877f.—Shells collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W.. 1: 311-321. , 1877g.—Continuation of the Mollusca of the Chevert Expedition. Proc. Linn. Soc. N.S.W., 1: 362-368. , 1877h.—Continuation of the Mollusca of the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 1-6. , 18771.—Continuation of the Mollusca collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 20-25. ————, 1877).—Continuation of the Mollusca collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 41-46. —— , 1877k.—Continuation of the Mollusca collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 46-53. , 1877/.—Continuation of the Mollusca collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 55-60. , 1877m.—Continuation of the Mollusca collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 74-89. , 1877n.—Desctiption of two new species of Helia, from New Guinea and the Louisiade Islands. Proc. Linn. Soc. N.S.W., 2: 120-122. = , 1878a.—Continuation of the Mollusca collected during the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 128-135. , 1878b.—Mollusca of the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 143-145. , 1878c.—Continuation of the Mollusca of the Chevert Expedition. Proc. Linn. Soc. N.S.W., 2: 368-369. — , 1878d.—Descriptions of seven new species of terrestrial and marine shells from Australia. Proc. Linn. Soc. N.S.W., 3: 77-81. , 1878e.—Mollusca of the Chevert. Expedition. Proc. Linn. Soc. N.S.W., 3: 155. = , 1879.—Desceription of a new species of Vivipara. Proc. Linn. Soc. N.S.W., 3: 221-222. —— , 1881.—Description of a new Bulimus from New Caledonia. Proc. Linn. Soc. N.S.W.. 6: 586-587. CiencH, W. J., and TuRNER, R. D., 1966.—Monograph of the genus Rhynchotrochus (Papuininae : Camaenidae). J. Malac. Soc. Aust., 1 (9): 59-87. , 1968.—Monograph of the genus Letitia (Papuininae: Camaenidae). J. Malac. Soc. Aust., 1 (11): 32-49. Cox, J. C., 1868.—A Monograph of Australian Land Shells. William Maddock, Sydney, 111 pp. ————, 1870.—Descriptions of seventeen new species of land shells from the South-Sea Islands, in the cabinet of Mr. John Brazier, of Sydney. Proc. Zool. Soc. (Lond.), 1870: 81-85. , 1892.—Descriptions of some new species of pulmonate Mollusca from Australia and the Solomon Islands. Proc. Linn. Soc. N.S.W., (n.s.), 6: 565-570. PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vou. 97, Part 1 W. F. PONDER AND P. J. STANBURY DD Futon, H., 1904.—A critical list of the Sphaerospira section of Thersites. J. Malac., 11: 2-11. GOLDMAN, J., Hinu, L., and SransBury, P. J., 1969.—Type specimens in the Macleay Museum, University of Sydney. 11. Amphibians and reptiles. Proc. Linn. Soc. N.S.W., 93: 427-438. GRIFFIN, D., and Stansury, P. J., 1970.—Type specimens in the Macleay Museum, University of Sydney. V. Decapod crustaceans. Proc. Linn. Soc. N.S.W., 95: 122-131. Haun, E., 1962.—A List of the Designated Type Specimens in the Macleay Museum, Insecta. Duplicated book, Macleay Museum, 184 pp. Heptey, C., 1891.—The land molluscan fauna of British New Guinea. Proc. Linn. Soc. N.S.W., (2), 6: 67-116. , 1899. The Mollusca of Funafuti. Part 1. Gasteropoda. Aust. Mus. Mem., 3: 395—488. , 1900.—Studies on Australian Mollusca. Part 2. Proc. Linn. Soc. N.S.W., 25: 495-513. , 1901a.—A revision of the types of the marine shells of the ‘‘ Chevert ”? Expedition. Rec. Aust. Mus., 4: 121-130. , 1901b.—Studies on Australian Mollusca. Part 3. Proc. Linn. Soc. N.S.W., 25: 721-732. , 1901c.—Studies on Australian Mollusca. Part 4. Proc. Linn. Soc. N.S.W., 25: 16-25. , 1902a.—Studies on Australian Mollusca. Part 5. Proc. Linn. Soc. N.S.W., 26: 700-708. , 1902b6.—Studies on Australian Mollusca. Part 6. Proc. Linn. Soc. N.S.W., 27: 7-29. , 1908.—Studies on Australian Mollusca. Part 10. Proc. Linn. Soc. N.S.W., 33: 456-489. , 1922.—A revision of the Australian Turridae. Rec. Aust. Mus., 13: 213-359. , 1924.—Some naticoids from Queensland. Rec. Aust. Mus., 14: 154-162. TREDALE, T., 1934.—The freshwater mussels of Australia. Aust. Zool., 8: 57—78. ———, 1939.—A review of the land Mollusca of Western Australia. J. Roy. Soc. W.A., 25: 1-88. , 1941.—A basic list of the land Mollusca of Papua. Aust. Zool., 10: 51-94. , 1943.—A basic list of the fresh water Mollusca of Australia. Awst. Zool., 10: 188-230. TREDALE, T., and Huu, A. F. B., 1926.—A monograph of the Australian loricates. Part 6. Aust. Zool., 4: 164-185. ; KosBett, W., 1903.—Agnatha Systematisches Conchylien-Cabinet von Martini und Chemnitz, ed Kuster, Bd. 1, Abth. 12B, 362 pp. Laseron, C. F., 1956.—The families Rissoinidae and Rissoidae (Mollusca) from the Solanderian and Dampierian Zoogeographical Provinces. Aust. J. Mar. Freshw. Res., 7: 384-484. , 1959.—The family Pyramidellidae (Mollusca) from northern Australia. Awst. J. Mar. Freshw. Res., 10: 177-267. Macmittan, D. S., 1956.—A Squatter Went to Sea. Currawong Publishing Co., Sydney. 165 pp. MacrpuHerson, J. H., and GABRIEL, C. J., 1962.—Marine Molluscs of Victoria. Melbourne Univ. Press. 475 pp. Piussry, H. A., 1889.—Trochidae, Stomatidae, Pleurotomariidae, Haliotidae. Manual of Conchology, 11, 519 pp. , 1893.—Acanthochitidae, Cryptoplacidae and appendix. Tectibranchiata. Manual of Conchology, 15, 436 pp. SHarp, B., and Pinssry, H. A., 1897.—Scaphopoda. Manual of Conchology, 17, 348 pp. Smirn, EK. A., 1889.—On the land-and-freshwater-shells of the Louisiade Archipelago. Ann. Mag. Nat. Hist., Ser. 6, 4: 199-209. Sotem, A., 1959.—Systematics and zoogeography of the land and freshwater Mollusca of the New Hebrides. Fieldiana: Zool., 43: 1-359. STanpury, P. J., 1968.—Type specimens in the Macleay Museum, University of Sydney. I. Fishes. Proc. Linn. Soc. N.S.W., 93: 203-210. , 1969a.—Type specimens in the Macleay Museum, University of Sydney. III. Birds Proc. Linn. Soc. N.S.W., 93: 458-461. , 1969b.—Type specimens in the Macleay Museum, University of Sydney. IV. Mammals. Proc. Linn. Soc. N.S.W., 93: 462-463. "TAPPARONE-CANEFRI, C., 1883.—Fauna malacologica della Nuova Guinea e delle isole adiacenti. Ann. Mus. Civ. Storia Nat., Genova, 19: 1-313. TEntson-Woops, J. E., 1881.—On some new marine Mollusca. Trans. Roy. Soc. Vict., 17: 80-83. Tryon, G. W., 1884.—Conidae, Pleurotomidae. Manual of Conchology, 6, 413 pp. PROCEEDINGS OF THE LINNEAN Socrety or New SoutH WatzEs, Vou. 97, Part | GENETIC BASES FOR RESISTANCE IN TWO COMMON WHEAT CULTIVARS TO STEM RUST STRAINS OF UNUSUAL AVIRULENCE! A. K. SANGHI? AND EH. P. BAKER [Accepted for publication 17th November 1971] Synopsis The genetic basis for resistance in two cultivars, Pusa and Mona, of common wheat (Triticum aestivum L.) to certain avirulent strains of stem rust was investigated. These cultivars are susceptible to field strains of wheat stem rust (Puccinia graminis var. tritici Eriks. and Henn.). The stem rust strains employed comprised rye stem rust (P. graminis var. secalis Eriks. and Henn.) and strains classified as wheat stem rust but possessing unusual genes for avirulence. These latter strains included sexual and somatic hybrids between wheat and rye stem rusts. Pusa and Mona were shown to possess four and three genes, respectively. From the segregation patterns in hybrids between the parents and pathological tests on lnes carrying single genes conditioning resistance two genes were found to be common to each cultivar. One was identified as Sr18 previously described and shown to be present in varieties throughout the world. A total of five distinct genes was identified in these cultivars. Observations on the behaviour of lines possessing isolated genes with that of the parents indicated that additive gene action was involved in conditioning the hypersensitive parental infection types. INTRODUCTION Cultivars of common wheat (Triticum aestivum L. em. Thell.) are in general resistant to rye stem rust (Puccinia graminis var. secalis Eriks. and Henn.) but comparatively little is known about the genetic basis of such resistance or of its relationship to that conditioning resistance to wheat stem rust (P. graminis var. trittc? Eriks. and Henn.). However, since certain wheat genotypes are at least moderately susceptible to particular collections of rye stem rust some progress has been made in analyses of these aspects. Further, firstly sexual hybrids (Johnson, 1949) and asexual hybrids (Watson and Luig, 1959; Bridgmon and Wilcoxon, 1959) between wheat and rye stem rusts have been produced in the laboratory. Normally such hybrids are intermediate in pathogenic character- istics between the parental cultures and are frequently avirulent on wheat genotypes recognized as susceptible to field strains of wheat stem rust. Secondly, certain exceptional] strains classified as forma specialis or variety triteci of P. graminis from pathogenic attributes on various diagnostic hosts are likewise avirulent on some wheat genotypes normally susceptible to wheat stem rust strains. These two groups of rust strains are considered to possess unusual genes for avirulence. In the present study the genetic bases for resistance in the wheat cultivars Pusa and Mona to a culture of rye stem rust and to cultures of wheat stem rust with unusual genes for avirulence were investigated. These two cultivars are recognized as lacking resistance genes to all isolates of P. graminis var. tritict collected in the field on wheat. REVIEW OF LITERATURE The host-pathogen relationships of genes for avirulence in certain strains of rust identical with, or closely related to, those used in the current studies have ' Adapted from part of a Ph.D. thesis submitted to the University of Sydney by the senior author. The financial assistance of the Wheat Industry Research Committee of New South Wales and the University of Sydney is gratefully acknowledged. ? Present address : Agricultural Experiment Station, University of Udaipur, Udaipur, India. PROCEEDINGS OF THE LINNEAN Society oF New SoutH WaAtess, Vout. 97. Part 1 A. K. SANGHI AND E. P. BAKER D7 been studied in a number of investigations. Loegering and Powers (1962) inoculated 20 common wheat genotypes with 108 F, cultures from a cross between culture 111-55A (race 111) and culture 36-55A (race 36) of P. graminis var. tritics and concluded that eight independent genes for pathogenicity were involved. Berg et al. (1963) studied the genetic basis for resistance to culture 111—55A in crosses between Little Club (susceptible) and Marquis and Kota (resistant), respectively. They reported that three dominant genes conditioned resistance in each resistant variety. Sheen and Snyder (1964), employing Marquis chromosome substitution lines in a Chinese Spring background, located one gene on chromosome 2A and a second gene on chromosome 3B in Marquis conferring resistance against race 111. The same authors (Sheen and Snyder, 1965), using Kenya 338 chromosome substitution lines, found that seven substitution lines differed from the susceptible Chinese Spring background in conditioning perceptible resistance to a culture of race 111. Rondon et al. (1966) found two and three independent genes in P.I. 94701 and Reliance, respectively, operative against race 111. Sears et al. (1957) reported a gene for resistance to a culture of race 111 on chromosome 7A of Chinese Spring and Thatcher. Williams and Gough (1965) reported four genes, of which three were identified as Sr7, Sri3d and Sri4, respectively, in Khapli operative against race 111. Gough and Williams (1963) reported that three incompletely dominant genes in each of the cultivars Acme and Mindum (both 7. turgidum 1.) conditioned resistance to culture 111-SS82. The investigations of Luig and Watson (1965) showed that Hureka (7. aestivum) possessed two independent genes, one Sr6 operative against rust strains 103—H-2 and 111—-E-2, and the other SrH/ operative against 103-—H-2; Sr6 is also known to condition resistance to certain field strains of P. graminis var. tritict. The same authors explained the resistance of Gabo to strains 103-—H—2 and 111—E-2 on the basis of a single gene, which they designated Sr@G2. A second gene SrG‘3 in this cultivar controlled resistance to culture A20 of P. graminis var. secalis. They reported also that Charter possessed these two genes. Baker et al. (1970) located SrG2, which was redesignated as Sr/S in the sequence of gene symbols used to catalogue stem rust resistance genes in wheat, on chromosome 7D and concluded that the gene was present in a number of wheat cultivars throughout the world. Sanghi and Luig (1971) found it necessary to invoke five genes to explain the pattern in inheritance of resistance exhibited by Mentana to cultures identical with those employed in the current investigations. The gene Sr8, effective against certain field strains of wheat stem rust, also operated against all six cultures utilized. Four genes were considered to be present in Yalta; the gene Sr1/, known to condition resistance to many field wheat stem rust strains, operated against only two of the cultures. MATERIALS AND METHODS A short description of each of the cultivars is as follows : Pusa W801 (W numbers refer to the Sydney University Wheat Accession Register) is an Indian selection from local collections. The spikes are apically tipped and the grains white, vitreous and plump. Mona W1168 is a selection made in Australia from a cross Plowman’s No. 3xCanberra. The spikes are apically tipped and grains white. Morocco W1103 is of unknown pedigree. The spikes are fully awned with pubescent glumes. The grains are pale red in colour. Purple Straw W1816 is of unknown pedigree. The spikes are lax, apically tip awned with glabrous glumes. The grains are pale red in colour. W2691 is a line developed at Sydney University specifically for susceptibility to P. graminis var. secalis and to strains of P. graminis var. tritici with unusual genes for avirulence. It originated from an F, plant of the cross Little Club x (Gabo? x Charter). PROCEEDINGS OF THE LINNEAN Society oF New SoutrH Wates, Vou. 97, Part 1 58 GENETIC BASIS FOR RUST RESISTANCE IN WHEAT All five cultivars are fully susceptible to Australian field strains of stem rust collected on wheat. The nomenclature of the wheat stem rust strains employed follows that adopted by Watson and Luig (1963, 1966). The six stem rust strains used in inheritance studies were described in detail by Sanghi and Luig (1971) and originally by Watson and Luig (1959), Luig and Watson (1965) and Baker et al. (1970). The origin and characteristics of each strain are hence only here briefly described. Accession 57241 is a field culture of P. graminis var. secalis. Cultures 103-H-2 (58-L-1) and M-10b (58—-L-—2) originated as somatic hybrids between stem rust strain NR-2 (P. graminis var. tritici) and 57241. Culture 111—E-2 (56-L-1) is a strain of P. graminis var. tritict to which few wheat cultivars are susceptible. It could have arisen from a Gross between P. graminis var. tritict and P. graminis var. secalis (Luig and Watson, 1965). Culture H-42 (64-L-1) is a sexual hybrid between P. graminis var. tritici strain 111—K-2 and P. graminis var. secalis culture H-34 and combines to some extent the pathogenic range of both parents. It is virulent on Black Winter rye. Morocco and W2691 are among the few wheat cultivars which are susceptible to H—42. It does not correspond to any race of wheat stem rust described by Stakman et al. (1962) from its infection types on the standard differential set. Culture 71-0 (63666) was collected on Agropyron scabrum L. in Queensland in 1963. Itis astrain of wheat stem rust non-pathogenic on wheat genotypes commonly susceptible in the field. Pots of seedlings were inoculated with a suspension of uredospores in ‘* Mobil Sol 100 ”, a low viscosity light paraffin oil, used as a spore carrier. A ‘‘ Desaga Spray Can” was used to apply the oil-rust suspension. Artificial moisture, in the form of a suspended fog, was created in a glasshouse for 12-15 hours during incubation. After inoculation the pots were placed on benches in well-lit glass- houses maintained at temperatures of 70-+-5° F. Notes on rust infection types. were taken 12-14 days after inoculation. Seedling infection types were based on the scale proposed by Stakman et al. (1962). TABLE 1 Infection Types Hxuhibited by Seedlings of Pusa, Mona, Susceptible Wheat Cultivars and F, Plants Inoculated with Six Strains of Stem Rust Rust Straim Wheat Cultivar or Hybrid 5724] 103—H-—2 M10—b 111—E-2 H-42 71-0 Pusa W801 .. me pen): 0; ;1—-2= 0; 0; 2 Mona W1168 oft 2 ¢ 2=n : ; 2n W2691 a: a so Bae 3-4 213¢e 3j aL 3-4 2n Morocco W1103 a el 2— 3+ 3 3 2 Purple Straw W1816 32 — = 3+ 72 =2— 2= 34+ W 2691 « Pusa 0; 0; 2 0: 0; 2 Pusa x W2691 0; 0: 2 0: 0: 2 Morocco « Pusa 0; 0: ee 0; 0; L= Pusa x Morocco 0; 0; st 0 0: B= Purple Straw x Pusa 0: 0: Dee 0 0: 2+ Pusa x Purple Straw 0; 0; 2+ 0 0: 2+ W 2691 x Mona JL 32 = 3n : 3n Mona x W2691 Be 22 = 3n Q 3n Morocco x Mona : a 3n 2—n Mona x Morocco ah :2= 3n 2—n Purple Straw x Mona es ; 3n 3n Mona x Purple Straw ;2= ; 3n 3n PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, VoL. 97, Part 1 A. K. SANGHI AND E. P. BAKER 59 The genetic analyses of the resistant parents were based on results from the F,, F,, F; and F, generations of crosses with a susceptible parent. In the case of cultures 103-—H-2, 111—E-2, H-42 and 57241, the susceptible parent was W2691. Morocco was employed for M10-b and Purple Straw in the case of 71-6 since W2691 exhibited only moderately susceptible (‘‘ 2{3c’) infection types to these rust cultures. EXPERIMENTAL RESULTS The infection types exhibited by parental and F, seedlings to the six rust cultures are set out in Table 1. Pusa W801 The resistant infection type (‘‘ 0;”’) of Pusa was dominant in F, tests with cultures 103—H-2, 111—E-2, H-42 and 57241. In tests with cultures M10-b and 71-0, in which Moroceo and Purple Straw were used as susceptible parents, the F, seedling infection types indicated partial dominance of the Pusa reaction. Inheritance of Resistance to Cultures 103—H-—2, 111—E-2 and H-42 The F, population of the cross W2691 x Pusa was derived from 11 F, plants. The progenies of different plants were, in general, tested with culture 103—H-2, 111—-H-2 or H-42. In certain instances the same plant progeny was tested with more than one culture. The data are presented in Table 2. In a total of 2,386 seedlings, 41 were susceptible and the results suggested the presence of three independent genes conditioning resistance in Pusa to each of the three cultures. Infection types ranged from “‘0;”’, “‘;”’, ‘‘2=” (classified as highly resistant), ew14+”, “2—”, “2—2” (resistant—moderately resistant) to “3”, “34” (susceptible). The data from different families were homogeneous (P=0-50-0-30). The segregation pattern further suggested that each gene conditioned a different, but resistant, infection type. TABLE 2 Number of F, Seedlings in Different Infection Type Classes in the Cross W2691 Pusa Tested with Three Stem Rust Cultures Infection Types Vie Rust - 63:1 12 Culture HR R—MR S HR R Value 0;,3,2 = ;14,2=,2—2 3,3-+ Total MR:S 103—H-2 Eke 612 53 11 676 0-018 0-90—0-80 111—E-2 a 878 116 19 1013 0-646 0-50—-0-30 H-42.. ae 629 57 11 697 0-001 0-98—0-95 Total... 2119 226 4] 2386 0-377 0-70—0-50 HR=highly resistant ; R—MR=resistant to moderately resistant ; S—susceptible. The breeding behaviour of 239 F, plants to culture 103-—H-—2, presented in Table 3, conformed statistically to an expected ratio of 37 homozygous resistant : 26 segregating: 1 homozygous susceptible line and supported the hypothesis proposed from F, results. Of the 86 segregating lines, 59 segregated in a ratio approximating 63 resistant : 1 susceptible or 15 resistant : 1 susceptible seedling. A population of 50 seedlings in each F, line is needed to distinguish a 3: 1 from a 15:1 segregation ratio at P>0-95. In practically all cases the number in each line exceeded 50, and these two segregation classes could thus be distin- guished. However, a population of 235 individuals is needed to distinguish a 15:1 from a 63:1 segregation ratio. Hence these classes were not separated PROCEEDINGS OF THE LINNEAN Socitery OF NEw SoutH WALES, Vou. 97, Part 1 60 GENETIC BASIS FOR RUST RESISTANCE IN WHEAT in the analysis because seedling numbers were insufficient to determine into which class certain segregating lines should have been placed. The number of such lines approximated the expected 20/64 of the F, populations. Seedlings in 27 lines segregated in a 3 resistant: 1 susceptible ratio ; of these, 12 segregated for 3 ‘““2—”:1° 3+”, 8for3" j14+ ":1“3+4” and 7 for 3 ‘2,24 ”:1 ‘*34 ” infection type. The number of lines in each group approximated the expected 2/64 of the population. These results confirmed that Pusa possesses three genes which condition ‘‘2=”, “314 ” and ‘‘ 2” infection types, respectively, “for resistance to culture 103-H-2. The first two genes are dominant, whilst that conditioning a ‘‘ 2” infection type is incompletely dominant, since heterozygous genotypes showed ‘‘ 2+ ” infection types. TABLE 3 Behaviour of F, Lines of the Cross W2691 x Pusa Tested with Culture 103-H—2 Number of Lines F, Infection Types Ratio Classification or Reactions Observed Expected Expected Homozygous resistant .. Os == eile 151 138-17 37 Segregating brea .. 63R:18 or 15R:1S 59 74-69 20 Segregating ee eS (2 —) eel (Ssaito) 12 7:47 2 Segregating ta .. 351+) : 1(3,3-+) 8. 4-47 2 Segregating : . 3(2,2-+-): 1(3,3-+) 7 7°47 2 Homozyeous susceptible Sa} 2 mdi ] Total 239 239-00 64 ¥Z=8-104; P=0-20-0-10. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y*=3-265 ; P=0-20-0-10. R=resistant. The results of F, tests with cultures 111—E—2 and H—42, presented in Tables 4 and 5, respectively, showed a similar pattern of segregation indicating the presence of three independent genes in Pusa for resistance to these cultures. The genetic relationship between genes operative against the three cultures was studied when the same 239 F, lines were inoculated separately with cultures 103-—H-2 and 111—E-2. In like manner 104 F, lines were employed in correlated tests involving cultures 103-H-2 and H-42. A P value of <0-001 was obtained in each case from chi-square tests for independent behaviour of F, lines. Four TABLE 4 Behaviour of F, Lines of the Cross W2691 x Pusa Tested with Culture 111—E-2 Number of Lines F, Infection Types Ratio Classification or Reactions Observed Expected Expected Homozygous resistant .. O22 — ee 147 138-17 37 Segregating if .. 63R:18 or 15R:18 63 74-69 20 Segregating Na .. 3(32=2—): 1(3,3-+) 12 7-47 2 Segregating ae .. 3(;1+) : 1(3,3+-) 8 7:47 2 Segregating : . 3(2,2+) :1(3,3+) U %:47 2 Homozygous susceptible Soe 2 3°73 1 Total .. se —_ 239 239-00 64 72=6-638; P=0-30-0-20. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y?=1-888 ; P=0-50—0-: 30. PROCEEDINGS OF THE LINNEAN Society or New SourH WALES. Vou. 97, Part 1 A. K. SANGHI AND E. P. BAKER 61 lines segregating against culture 111—EK-2 were homozygous resistant to cultures 102—H—2 and two lines segregating in tests with H—42 were homozygous resistant when tested with 103-H-2. This discrepancy was probably due to misclassi- fication of lines with smaller seedling numbers. The high correlation observed suggested that the three genes conditioning resistance to each of the cultures 103-—H-2, 111-EK-2 and H-42, respectively, were identical. TABLE 5 Behaviour of F, Lines of the Cross W2691x Pusa Tested with Culture H—42 Number of Lines FE Infection Types Ratio Classification or Reactions Observed Expected Expected Homozygous resistant .. 0;,;2=,2—,;1+.,2 208 196-56 37 Segregating of .. 63R:1S or 15R:18 91 106-25 20 Segregating oe .. 3(62=) :1(3,3+) 16 10-63 2 Segregating A .. 3831+) :1(3,3+) 12 10-63 2 Segregating ae .. 3(2,2+):1(3,3+) 9 10:63 2 Homozygous susceptible 3,3-++ 4 5-31 1 Total... zh — 340 340-01 64 x2=6-324; P=0-30-0-20. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y2=1-212 ; P=0-70—-0-50. Inheritance of Resistance to Culture 57241 of Rye Stem Rust The F, generation of the cross W2691 x Pusa consisted of 577 seedlings derived from three F, plants. A chi-square test for heterogeneity indicated that the data from different families were homogeneous (P=0-70-0-50) and the results from the families were therefore combined. In a total of 577 seedlings, 530 were highly resistant (‘‘ 0’, ‘‘ 1’, ‘‘2= ” infection types), 37 were resistant (3142, “2—”, “2—” infection types), and 10 were moderately susceptible (‘‘2+3c” infection types). These data conformed to a segregation ratio of 63 resistant: 1 moderately susceptible seedling and suggested the presence of three independent genes in Pusa conditioning resistance to this culture. Tests involving 228 F, lines (Table 6) confirmed the hypothesis proposed on the basis of F, tests. According to the hypothesis 37/64 would be homozygous resistant, 26/64 segregating and 1/64 homozygous for moderate susceptibility. Among the TABLE 6 Behaviour of F, Lines of the Cross W2691 x Pusa Tested with Culture 57241 Number of Lines F; Infection Types Ratio Classification or Reactions Observed Expected Expected Homozygous resistant .. 0;,;,2 =,;1+:12—,2 134 131-81 37 Segregating oe .. 63R:1MS or 15R:1MS 78 71-25 20 Segregating 70 .. 3(31+) : 1(2$3c) 6 Ue 133 2 Segregating ~ .. 3(2—,2+) :1(213c) 7 7°13 2 Segregating a .- 3(;12—,X —) : 1(2]38c) 4 9 183 2 Homozygous moderately susceptible a oN 213¢ 2 3°56 1 Total .. 38 —- 228 228-01 64 MS=moderately susceptible. 7z=2-459; P=0-80-0-70. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y?=0-726 ; P=—0-70—--050. PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SourH WALES, Vou. 97, Part 1 62 GENETIC BASIS FOR RUST RESISTANCE IN WHEAT segregating lines 20/64 would segregate for 63 resistant : 1 moderately susceptible or 15 resistant : 1 moderately susceptible and 6/64 for 3 resistant : 1 moderately susceptible seedling. In the latter category three groups of infection types were distinguished in populations derived from 17 F, plants. These groups were characterized by ‘‘;51+”, “2—” and ‘12—” seedling infection types, respectively, on homozygous genotypes. When tested statistically the number of lines observed in each of these categories did not deviate significantly from the number expected. The F, data, when grouped, conformed to a ratio of 37 homozygous resistant : 26 segregating : 1 homozygous for moderate susceptibility and indicated the presence of three independent genes conditioning resistance to culture 57241. To study the genetic relationship between genes for resistance in Pusa to cultures 103-—H-2, 111—E-2, H—42 and 57241, 79 resistant and four susceptible lines from the cross W2691 x Pusa, of which the reactions to cultures 103—H-2, 111-E-2 and H-42 were known, were selected and inoculated with 57241. Homozygous resistant lines with ‘“‘;1+ ” and “2” infection types to 103—H-2, 111—-E-2 and H-42 were also homozygous resistant to 57241 and exhibited similar infection types. One homozygous line showing ‘‘2—” to 103-H-2, “2=2= ” to 111—E-2 and “ ;2= ” infection types to H—42 exhibited ‘‘ 213c ” infection types to 57241, whilst two lines homozygous susceptible to 103—H-2, 111-E-2 and H-42 gave ‘‘;12—” infection types to 57241. These results indicated that the gene conditioning a ““2=” to 103-H-2, a “ ;2=2—” to 111—-E-2 and a “ ;2= ” infection type to H—42 is not operative against 57241 and that the gene conditioning a ‘‘ ;12— ” infection type to 57421 is ineffective against cultures 103-—H-2, 111—E-2 and H-42. This infers that Pusa has a fourth gene for resistance. The dominant gene giving a ‘‘2=—” to 103-H-2, ““.2=2—=” to 111-H-2 and a ‘“‘;2=” infection type to H—42 is inoperative against 57241. The second and third genes condition ‘‘ ;1-+ ”’ and “‘ 2” infection types, respectively, against all four cultures ; the second behaves in a dominant manner, whilst the third is partially dominant and when heterozygous conditions a “2+” infection type with all four cultures. The fourth gene conditions a ‘* -12— ” infection type and operates against 57241 alone. This gene is partially dominant and when heterozygous conditions an ‘“‘ X — ” infection type. These four genes are designated provisionally SrPsi, SrPs2, SrPs3 and SrPs4, respectively. The first gene does not operate against culture 57241, the fourth does not operate against cultures 103-—H-2, 111—H-2 and H-42, while the second and third operate against all four cultures. Inheritance of Resistance to Culture M10-b The results of F, seedling tests in the cross Morocco (susceptible) x Pusa (resistant) indicated the presence of two independent genes for resistance in Pusa to this culture. In a total of 660 F, seedlings derived from three F, plants, 612 were resistant to moderately resistant (‘ ;12= ”—‘‘2+ ” infection types) and 48 were susceptible (‘‘ 3,3-+ ” infection types). This number conformed to a 15 resistant : 1 susceptible seedling ratio (P=0-30—-0-20). The results (Table 7) of the progeny tests of F, plants agreed with the ratio of 7 homozygous resistant: 8 segregating: 1 homozygous susceptible F, line expected on this hypothesis. Further evidence for the presence of two genes was obtained from progeny tests of F, plants of the cross Purple Straw x Pusa. Purple Straw, susceptible to culture M10-b in crosses with Pusa, also gave a two-gene segregation pattern in F, (Table 7). Individual F, lines segregating monofactorially gave a 1 resistant : 2 intermediate : 1 susceptible rather than a 3 resistant: 1 susceptible seedling segregation ratio, indicating that both genes for resistance in Pusa against culture M10-b are partially, rather than completely, dominant. PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SourH Wates, Vou. 97, Part 1 A. K. SANGHI AND HE. P. BAKER 63. Inheritance of Resistance to Culture 71-0 The F, populations from reciprocal crosses between Purple Straw and Pusa totalled 468 plants derived from five F, plants. A chi-square test on combined data for a 15 resistant : 1 susceptible seedling ratio gave a P value of 0-30-0-20. This indicated the presence of two independent genes in Pusa operative against 71-0. This was confirmed in F, tests. The breeding behaviour of F, plant progenies agreed with an expected ratio of 7 homozygous resistant : 8 segregating : 1 homozygous susceptible F, line. Fifty-one F, lines from the cross Purple Straw x Pusa were studied to investigate the relationship between genes operative against cultures M10-b and 71-0. An identical breeding behaviour was obtained, indicating that the same genes operate against both cultures. TABLE 7 Behaviour of F', Lines of the Crosses Morocco x Pusa and Purple Straw x Pusa Tested with Cultures M10-b and 71-0 Rust 2 1p Cross Culture R Seg Ss Total 788 ll Value Morocco x Pusa M10-b 30 37 5 72 0-156 0-95-0-90 Purple Straw < Pusa .. .. M10—b 18 28 5 51 2-109 0-50-0-30 Purple Straw x Pusa .. 22 @1=0 18 28 5 51 2-109 0-50—-0:30 Total od — 66 93 15 174 3:325 0-20-0-10 Seg =segregating. Isolation of Genes Fifty-seven F, lines from the cross W2691 x Pusa were tested to study the relationship between the genes operative against the six rust cultures. These F, lines were derived from eight F, lines segregating monogenically for 3 C=?) al (6 BALM), 3 (C2 i SL) 2 FL (G8 BL and 3 (GEROE 2 Vicon (EBA ») infection types, respectively, and two F’, lines homozygous susceptible when tested with culture 103—H-2. Seedlings of separate sowings of each line were tested with each of the six cultures simultaneously in the same glasshouse. The results supported the hypothesis that each selected F, line carried a single gene for resistance and confirmed that the two susceptible lines were homozygous for susceptibility to culture 103—H-2 ; F, lines homozygous for resistance to culture 103—H-2 due to the gene SrPs1 (conditioning a ‘‘2=” infection type) were homozygous resistant to cultures 111—E-2 and H-42 (“ ;2=2= ” and“ ;2=” infection types, respectively) but were homozygous for moderate susceptibility to cultures 57241, M10-b and 71-0. These results indicated that SrPs/ is ineffective against the latter three cultures and that moderate susceptibility in these lines is due to the W2691 genotype. Lines homozygous for resistance to culture 103—H-? due to the gene SrPs2 (conditioning a “ ;1+ ” infection type) were homozygous for resistance to cultures 111-E—2, H—42 and 57241 but homozygous for moderate susceptibility to cultures M-10b and 71-0. Lines homozygous for resistance to culture 103-H-2 due to SrPs3 (conditioning a ‘‘2” infection type) were homozygous resistant to all six cultures. Two F, lines, which were homozygous susceptible to cultures 103-H-2, 111-E-2 and H-42, were resistant to cultures 57241, M10—b and 71-0 and exhibited ‘‘;12—” infection types to these cultures. These tests indicated that these lines did not carry any of the abovementioned genes and that resistance was apparently due to the fourth gene SrPs4. PROCEEDINGS OF THE LINNEAN SOCIETY oF NEw SoutH WALES, Vou. 97, Part I 64 GENETIC BASIS FOR RUST RESISTANCE IN WHEAT SrPs3, one of the two genes operating against cultures M—10b and 71-0, operates against all cultures used and SrPs4, the second gene operating against cultures M10-—b and 71-0, provides resistance against culture 57241 but not cultures 103—H-2, 111—EK-—2 and H-42. The results obtained from the genetic analysis of Pusa indicated that it possesses a total of four genes for resistance to the six stem rust cultures utilized. The infection types on seedlings of selected F, lines with different homo- zygous genotypes for resistance are presented in Table 8. TABLE 8 Seedling Infection Types Exhibited by Parents and Selected F', Lines of the Cross W2691« Pusa Tested with Sia Stem Rust Cultures Rust Culture Effective Pedigree Gene(s) 103—H-2 111—H-2 H-42 57241 M10—b 71—0 1165.62.2.87.4 i y= 2=2— 2= 2T3¢ 2t3e 2t3e SrPsl 1165.62.2.34.4 epee ;1+ sL+ ie 2t3e 213e SrPs2 1165.62.2.133.1 2 2 2 2— 2 Y SrPs3 1165.62:2.104.1 .. 3 3 3 si(2j3— gilD— 12) — SrPs4 Pusa a Me O; O: QO; Q; -] —2— — SrPsl SrPs2 SrPs3 SrPsd W 2691 3+ 3 3-4 2T3¢ 213¢ 2T3E — Mona W1168 The resistant infection type of Mona was dominant in the case of cultures 103-—H-2, 111—E-2 and H-—42 but incompletely dominant with cultures 57241, M10-b and 71-0 in F, seedling tests (Table 1). Inheritance of Resistance to Cultures 103—H-2, 111—E-2 and H-42 When tested with cultures 103—H-2, 111—E-—2 and H-42 the F, populations segregated into a ratio of 63 resistant : 1 susceptible seedling (Table 9). Thus the action of three independent genes for resistance in each case was inferred and this hypothesis was confirmed in F, tests. TABLE 9 Number of F, Seedlings in Different Infection Type Classes in the Cross W2691 x Mona Tested with Three Stem Rust Cultures Infection Types Ne Cross Rust = Total 63:1 12 Culture HR—R R Ss HR—R Value Q=2=9— 29 9 Bh +R:S W2691 «Mona 103—H—2 546 60 14 620 1-950 0-20-0-10 111—K-2 1084 38 16 1138 0-181 0-70—0-50 H—42 1083 50 10 1143 3:513 0:50-0:10 Total DNF 148 40 2901 0-636 0:50-0:30 The breeding behaviour of progenies of F, plants agreed with the expected ratio of 37 homozygous resistant : 26 segregating : 1 susceptible F, line when tested with each of the three cultures. Two hundred and thirty-four F, lines were tested with culture 103—H—2 (Table 10). On the basis of the hypothesis proposed, 37/64 of the lines would be homozygous resistant, 20/64 would segregate in a ratio of 63 resistant : 1 susceptible or 15 resistant : 1 susceptible, 6/64 would segregate in a ratio of 3 resistant: 1 susceptible seedling and 1/64 would be PROCEEDINGS OF THE LINNEAN Socrmpty or New SourH Wates, Vou. 97, Part 1 A. K. SANGHI AND E. P. BAKER 65 homozygous susceptible. When tested statistically the number of lines observed in each of these categories did not deviate significantly from that expected. TABLE 10 Behaviour of F, Lines of the Cross W2691 x Mona Tested with Culture 103—H-2 F, Number of Lines Ratio Classification Infection Types Expected Observed Expected Homozygous resistant a) 144 135-28 37 Segregating oh .. 63R:1S or 15R:18 62 73-12 20 Segregating ss A 3(2=1) :1(3,3+) 16 14-62 4 Segregating ae as 3(2,2+) :1(3,3+) 6 FoBil 2 Homozygous susceptible 3,30 6 367 1 Total .. ce _ 234 234-00 64 7¥i=4-122; P=0-50-0-30. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y?=2-885 ; P=0-30—0-20. A total of 291 F, lines was tested with culture 111-E-2 (Table 11). Five were homozygous for ‘‘ 3,3-+ ” infection types. This proportion of susceptible lines (approximately 1/64 of the population) supported the hypothesis that Mona possesses three independent genes for resistance. This was indicated further by the occurrence of 33 lines (approximately 6/64 of the population) which segregated in a 3 resistant: 1 susceptible seedling ratio. Occasionally within this group, individual progenies segregated in a 1 (“2”):2 (“24 ”):1 (“3” infection types) seedling ratio. The range of infection types among resistant segregants thus indicated incomplete dominance of one gene. TABLE 11 Behaviour of Ff, Lines of the Cross W2691< Mona Tested with Culture 111—H-2 F, Infection Types Number of Lines Ratio Classification or Reactions Expected Observed Expected Homozygous resistant eho, 185 168-23 37 Segregating es .. 63R:1S or 15R:18 68 90-94 20 Segregating .. .. 3&32=2=) : 1(3,3+) 8 9-09 2 Segregating ee. co) (ASHI) : 1(3,3-+) 13 9-09 2 Segregating .. .. 3(2,2+) :1(3,3+) 12 9-09 2 Homozygous susceptible 3,39-+ 5 4-55 1 Total .. 3 — 291 290-99 64 7¥Z=10-201; P=0-10-0-05. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y2=4-222 ; P=—0-20-0-10. Three groups of resistant infection types were exbibited among lines segregating 3 resistant : 1 susceptible seedling. The number of lines comprising each group was close to 2/64 of the F,; populations tested. These groups were characterized by “ ;2=2—”, “2=n” and “2” resistant infection types, respectively. Asin the case of Pusa crosses, no attempt was made to discriminate between lines segregating 63 resistant : 1 susceptible or 15 resistant : 1 suscertible seedling. The proportion of lines in this combined segregating class approximated the expected 20/64. Results from tests of 152 F, lines with culture H—42 (Table 12) were similar to those described for culture 111-E-2. Three lines were homozygous for PROCEEDINGS OF THE LINNEAN Society oF New SoutH Watzs, Vou. 97, Part ] 66 GENETIC BASIS FOR RUST RESISTANCE IN WHEAT ‘““3,3-+ ” infection types. The number of lines in each category which segregated for 3 resistant: 1 susceptible seedling was close to that expected for a three independent gene segregation. TaBLE 12 Behaviour of F, Lines of the Cross W2691 Mona Tested with Culture H—-42 F, Infection Types Number of Lines Ratio Classification or Reactions Expected Observed Expected Homozygous resistant 32=,2—,2—,2 93 87-88 37 Segregating he 2.) ©663RzIIS or VRS 40 47-50 20 Segregating ais oa 3(;2=) :1(3,38-+) 7 4-75 2 Segregating ie alo 3(2=n) : 1(3,3+) 4 4-75 2 Segregating te “fe 3(2,2+) : 1(3,3-+) 5 4-75 2 Homozygous susceptible 3,3 3 2-38 I Total . . Ae — 152 152-01 64 1z2—3°600; P=0-70-0-50. For an expected 37 resistant : 26 segregating : 1 susceptible ratio, y>=0-999 ; P=0-70-0-50. One hundred and fifty-two F, lines from the cross W2691 x Mona were used to study the relationship between genes conferring resistance to cultures 103—H-2, 111-E-2 and H-42. A high positive correlation was observed for behaviour of F, lines to the three cultures indicating that the same genes were operative against cultures 103-—H—2, 111-E-2 and H-42. These factors are designated tentatively Sr MnJ conditioning a ‘‘ 2=— ” infection type to 103—H-2, a ‘‘2=2= ” infection type to 111—-EK-2 and a “;2=” infection type to H-—42, SrMn2 conditioning a ‘‘2—n”’ infection type and SrMn3 conditioning a “ 2°’ infection type to all three cultures. Inheritance of Resistance to Culture 57241 of Rye Stem Rust A total of 573 F, seedlings derived from three F, plants was tested with culture 57241. The plants in each family segregated into three groups of infection types, “‘2=2= ”, ‘““2—2” and ‘‘ 213c”’, respectively. The chi-square for heterogeneity indicated that segregation in the individual families was similar (P=0-80-0-70) and the data from all families were therefore combined. The F, data, which conformed to a segregation ratio of 13 resistant (“‘2=2=— ”, ‘“2—2):3 moderately susceptible (‘‘213c”’ infection types) seedlings (P=0-80-0-70), indicated the presence of an incompletely dominant and a recessive gene for resistance in Mona to this culture. Tests in F, confirmed the hypothesis proposed and progenies of F, plants conformed statistically to a ratio of 7 homozygous resistant : 8 segregating : 1 line homozygous for moderate susceptibility ; of the 181 lines tested, 69 were homozygous resistant, 99 segregated and 13 were homozygous susceptible. There was no apparent difference in results from reciprocal crosses. One hundred and nine F, lines from the cross W2691 x Mona were tested to study the relationship between genes operative against cultures 103—H-—2 and 57241. Seedlings of separate sowings of each line were tested with each of the two cultures simultaneously in the same glasshouse and the results are presented in Table 13. A P value of <0-001 was obtained for a chi-square test for independent behaviour of F, lines to the two rust cultures. These results indicated that at least one gene is effective against both cultures. Of the 109 lines tested, 65 were homozygous resistant (‘ ;2=°-—‘‘ 2” infection types) to culture 103—H-—2 and 56 to culture 57241. Of the 65 lines homozygous resistant to culture 103-H-2, 45 were homozygous resistant, 10 segregated and 1 was PROCEEDINGS OF THE LINNEAN SociETy oF NEw SoutH WALES, Vou. 97, Part 1 A. K. SANGHI AND HE. P. BAKER 67 homozygous for moderate susceptibility (‘‘ 2t3c’ infection types) in tests with 57241. The line moderately susceptible to culture 57241 exhibited ‘ 2= ” infection types to 103—H-2 indicating that it possessed Sr Mn, which is therefore not operative against 57241. The behaviour of lines carrying the second gene designated SrMn2, which gave a ‘‘2=n” infection type to both cultures, indicated that this gene was recessive in tests with 57241 but dominant in the case of 103—H-2. The lines carrying the third gene designated Sr Mn3 exhibited ‘‘ 2” infection types to both cultures. This gene behaved in a partially dominant fashion conditioning a ‘*‘ 2+ ” infection type when heterozygous. These results indicated that the two genes giving resistance to culture 57241 also operate against 103-H-2. TABLE 13 Behaviour of F, Lines of the Cross W2691x Mona Tested with Cultures 103-H—2 and 57241 Behaviour Behaviour to Culture 57241 to Culture Total 103—H-2 Moderately Resistant Segregating Susceptible Resistant .. age 45 19 1 65 Segregating a 11 29 2 42 Susceptible ae = 1 1 2 Mowe 5 56 49 4 109 For independence, y3=36-856; P= <0-001. Inheritance of Resistance to Culture M10-b Four hundred and forty-seven F, generation seedlings tested with culture M10-b segregated into two infection type classes—‘‘ 2=n ”’—‘‘ 3n ”’ (375 seedlings) and ‘“ 3,3-+ °’ (72 seedlings). These numbers of plants in the two classes gave a satisfactory fit to a ratio of 13 resistant : 3 susceptible (P=0-20—0-10), indicating the presence of two independent genes for resistance in Mona, one dominant and the other recessive. This hypothesis was confirmed in F, tests (Table 14). A P value of 0:80-0:70 was obtained for an expected ratio of 7 homozygous resistant : 8 segregating: 1 homozygous susceptible line in the cross between Mona and Morocco. Evidence for the action of one incompletely dominant and one recessive gene was further obtained from progeny tests of F, lines of the eross Purple Straw x Mona tested with M10-b. A P value of 0:30-0:20 was obtained for an expected ratio of 7 homozygous resistant : 8 segregating : 1 homo- zygous susceptible line. TaBLE 14 Behaviour of F', Lines of the Cross Mona x Morocco Tested with Culture M10-b and Purple Straw Mona Tested with Cultures M10-b and 71-0 Rust Ratio 12 Cross Culture R Seg S Total Ex- Mie Value pected Mona x Morocco .. M10—-b 36 46 4 86 7sSsil — Oc@sl Ossveoew Purple Straw x Mona M10-—b 27 44 7 78 TeSBell BeOay Oesl=0cex Purple Straw x Mona 71-0 16 39 23 78 ie 2g il Woz Oxgdo0-a0) Inheritance of Resistance to Culture 71-0 Four hundred and nine F, plants tested with culture 71-0 were derived from three F, plants. A chi-square test for heterogeneity indicated that the PROCEEDINGS OF THE LINNEAN Society or New SoutH Watszs, VoL. 97, Part [| 68 GENETIC BASIS FOR RUST RESISTANCE IN WHEAT data from different families were homogeneous (P=0-70—0-50) and the data were combined. Based on infection types, the plants were classified into two groups in an observed ratio of 300 with ‘‘ 2-3n ”’ and 109 with “3,34” infection types. The observed segregation was an acceptable fit to a 3 resistant: 1 susceptible seedling ratio. These results were therefore best explained on the basis of a single gene difference for infection type. This hypothesis was confirmed in F, tests (Table 14). Seventy-eight F, lines segregated in 1 homozygous resistant : 2 segregating : 1 homozygous susceptible seedling ratio. These 78 F, lines were used in studying the relationship between genes conditioning resistance to cultures M10-—b and 71-0. The data, presented in Table 15, showed a P value of <0-001 for a test of independent behaviour, and indicated that genes in common were involved in the two cases. Twenty-seven lines were resistant to culture M10—b and 16 to 71-0. Five lines, which were resistant to culture M10—-b (exhibiting ‘‘ 2= ” infection types), were susceptible to culture 71-0 but no line susceptible to M10—b was resistant to 71-0. Among the 44 lines segregating to culture M10—b, 11 were susceptible but none was homozygous resistant to 71-0. These results indicated that the genes con- ditioning ‘‘ 2” infection types to M10-b and 71-0 are identical and that the gene conditioning a ‘‘2—n”’ infection type to M10—b did not operate against culture 71-0. TABLE 15 Correlated Behaviour of Seedlings of F., Lines of the Cross Purple Straw x Mona Tested with Cultures M10-b and 71-0 Behaviour Behaviour to Culture M10—-b to Culture Total 71-0 Resistant Segregating Susceptible Resistant .. oe 16 —- — 16 Segregating 1 6 33 = 39 Susceptible ies 5 11 u 23 Motalaer 27 44 7 78 For independence, y3=57-603; P value= <0-001. Isolation of Genes Thirty-two F, lines from the cross W2691 x Mona were tested to study the relationship between the genes operative against the six rust cultures. These F, lines were derived from six lines segregating monogenically for ‘“‘2= ”, ‘“2-n” and “ 2,2+ ” infection types, respectively, when tested with culture H-42. Seedlings of separate sowings of each line were tested with each of the six cultures simultaneously in the same glasshouse. The results supported the hypothesis that each selected F; line carried a single gene for resistance to culture H-42; F, lines homozygous for resistance to culture H-42 carrying SrMnI (‘‘;2— ” infection type) were also homozygous for resistance to 103-H-2 and 111—K-2 but homozygous for moderate susceptibility to cultures 57241, M10-b and 71-0. These tests inferred that the gene Sr Mn is not operative against the latter three cultures and that the moderate susceptibility observed was due to the W2691 genotype which exhibits a ‘‘2{3e ” infection type to these cultures. Each F, line homozygous resistant to culture H—42 due to the action of SrMn2 (‘“2—=n” infection type) were also homozygous resistant to cultures 103—H-2. 111—E-2, 57241 and M10-b but were homozygous for moderate susceptibility to culture 71-0. This gene behaved in a dominant manner with 103-H-2, 111—E-—2 and H-—42 but was recessive with cultures 57241 and M10-b. Lines homozygous in F, for resistance to culture H—42 due to the presence of SrMn3 (‘2 ” infection type) were homozygous resistant to all cultures. PROCEEDINGS OF THE LINNEAN SociETy or NEw SoutH WALES, Vou. 97, Part 1 A. kK. SANGHI AND Hi. P. BAKER 69 Tests with the six rust cultures thus revealed that Mona has three genes conditioning resistance to the cultures employed. The seedling infection types of selected F, lines with different homozygous genotypes for resistance are presented in Table 16. TABLE 16 Seedling Infection Types Exhibited by Parents and Selected Ff, Lines of the Cross W2691~% Mona Tested with Sia Stem Rust Cultures Rust Culture Effective Pedigree Gene(s) 1165.66.7.25.1 .. 2= 2=2— 32= 213¢ 243¢ 243¢ SrMni 1165.66.7.53.3 .. 2=n =n 2=n 2=n =i 2438e SrMn2 1165.66.7.133.1 D 2 2 2 — 2 Qn SrMn3 Mona . ie ;2= 5 : 2 =2 = 2—=n 2n SrMn1 SrMn2 SrMn3 W2691 .. she Ae epee Sj 3-4 213c 243c 2138¢ = DISCUSSION A number of genes in hexaploid wheat specifically conditioning resistance to cultures of P. graminis var. tritici possessing unusual genes for avirulence, P. graminis var. secalis and sexual or somatic hybrids between these formae speciales were revealed in the present study. A total of five genes in this category was found in two cultivars, neither of which is known to possess resistance to Australian field strains of wheat stem rust. The cultivar Pusa was shown to possess four, and Mona three, genes, respectively, conditioning resistance. On the bases of information from both genetical and pathological tests the genes SrPsi and SrMn1, on the one hand, and SrPs3 and SrMn3, on the other, are allelic. Evidence in this connection was based on the absence of segregation in F, and F, generations in crosses between the parental genotypes (Sanghi, 1968). The gene SrPs/ is the same as reported by Luig and Watson (1965) in Gabo and Charter, Williams e¢é¢ al. (1966) in Marquis and Reliance, Sears et al. (1957) in Hope, Sanghi (1968) and Baker et al. (1970) in Purple Straw and Mentana. Kaveh et al. (1968) found that the gene designated Mq’A in Marquis was allelic, or very closely linked, with a gene for rust resistance on chromosome 1D of Hope. By aneuploid analysis Anderson and Williams (1968) placed Mq.A in Marquis and A#l.A in Reliance on chromosome 1D. Baker ef al. (1970) designated their gene Sri8 and located it on chromosome 1D by aneuploid analysis. They also concluded that Sr18 is widespread in wheat cultivars. Because of the presence of this gene in various unrelated cultivars, it is difficult to trace its origin. No relationship between SrPs2, SrPs3, SrPs4 and SrMn2 and other previously reported genes could be established. A study of segregation in crosses between lines carrying isolated genes for resistance in an appropriate common background is needed to establish genetic relationships in such instances. The gene SrMn2 in Mona was dominant in tests with cultures 103-H-2, 111—-H-2 and H—42, but recessive to cultures 57241 and M10-b. The phenomenon of reversal of dominance of resistance to rust has been reported by Knott and Anderson (1956), who noted that the gene Sr6 was dominant to race 56 but recessive in the case of 15B. Despite the presence of multigenic systems in both parents conditioning resistance to the specific rust cultures employed, it was possible to isolate singly the host genes involved. Observations on the behaviour of lines of this type in PROCEEDINGS OF THE LINNEAN SocteTY oF NEw SoutH WALES, Vou. 97, Part 1 70 GENETIC BASIS FOR RUST RESISTANCH IN WHEAT comparison with the parental reactions afford opportunities for studying the nature of gene interaction in conditioning rust infection types. Additive gene action was indicated in the present instance. For example, the hypersensitive ‘0; infection type exhibited by Pusa to culture 103-H-—2 was due to the additive effects of genes SrPs1, SrPs2 and SrPs5, which individually condition “2=”,**;1+ ” and “ 2 ” infection types, respectively. SUMMARY The genetic basis for resistance to certain stem rust strains possessing unusual genes for avirulence was studied in two wheat parents Pusa and Mona, susceptible to field strains of wheat stem rust. These rusts included two cultures of wheat stem rust, one (race 71-0) collected on Agropyron scabrum in the field and the other (race 111—E-2) presumably originating from barberry infection. In addition one culture of rye stem rust (culture 57241) and certain sexual (culture H-42) or somatic hybrids (cultures 103-—H-—2 and M10-b) between wheat and rye stem rusts were employed. Tests involving F,, F,, F, and F', generations from the crosses of Pusa and Mona (resistant) with susceptible parents indicated that the resistance in the case of the former was conditioned by four genes and in the case of the latter by three genes. The relationship between certain of these genes was established. From the segregation patterns in hybrids between the parents and pathological tests on isolated lines carrying single genes conditioning resistance SrPsi (Pusa) and SrMn1 (Mona), on the one hand, and SrPs3 and SrMn3, on the other, were allelic. A total of five genes in both parents were identified. The genes SrPs1 and SrMn1 were considered to be idential with Sr/6, previously described and shown to be present in varieties throughout the world. References AnpERSON, M. K., and Wittiams, N. D., 1968.—Aneuploid analysis of genes for stem rust resistance derived from Marquis and Reliance wheat. Agron. Absir., 1968: 2. Baker, HE. P., Saneur, A. K., McIntosu, R. A., and Lute, N. H., 1970.—Cytogenetical studies in wheat. III. Studies of a gene conditioning resistance to stem rust strains with unusual genes for avirulence. Aust. J. biol. Sci., 23: 369-375. Bere, L. A., Goucu, F. J., and Wiiuiams, N. D., 1963.—Inheritance of stem rust resistance in two wheat varieties Marquis and Kota. Phytopathology, 53: 904-908. Bripemon, G. H., and Witcoxon, R. D., 1959.—New races from mixtures of urediospores of varieties of Puccinia graminis. Phytopathology, 49: 428-429. Goueu, F. J., and Wriiiiams, N. D., 1963.—Inheritance of stem rust reaction in two durum varieties Acme and Mindum. Phytopathology, 53: 295-299. JOHNSON, T., 1949.—Intervarietal crosses in Puccinia graminis. Can. J. Research (C), 27: 45-65. Kaveu, H., Wituiams, N. D., and Goueu, F. J., 1968.—Allelic and linkage relations among genes for reaction to wheat stem rust. Agron. Abstr., 1968: 12. Knott, D. R., and AnpERSoN, R. G., 1956.—The inheritance of rust resistance. J. The inheritance of rust resistance in ten varieties of common wheat. Can. J. agr. Sci., 36: 174-195. LorGERiING, W. Q., and Powers, H., Jr., 1962.—Inheritance of pathogenicity in a cross of physio- logical races 111 and 36 of Puccinia graminis f. sp. tritici. Phytopathology, 52: 547-554. Luic, N. H., and Warson, I. A., 1965.—Studies on the genetic nature of resistance to Puccinia graminis var. tritici in six varieties of common wheat. Proc. Linn. Soc. N.S.W., 90: 299-327. Ronpon, M. R., Gouen, F. J., and Witiiams, N. D., 1966.—Inheritance of stem rust resistance in Triticum aestivum ssp. vulgare ““ Reliance’ and P.I. 94707 of Triticum durum. Crop Scv., 6: 177-179. Saneut, A. K., 1968.—Studies on the genetic nature of resistance in common wheat to straims of stem rust possessing unusual genes for avirulence. Ph.D. thesis, Univ. of Sydney. Saneut, A. K., and Lure, N. H., 1971.—Resistance in wheat to formae speciales tritici and secalis of Puccinia graminis. Can. J. Genet. Cytol. (in press). Sears, HE. R., Lozrcerine, W. Q., and RoDENHISER, H. A., 1957.—Identification of chromosomes carrying genes for stem rust resistance in four varieties of wheat. Agron. J., 49: 208-212. PROCEEDINGS OF THE LINNEAN Society or New Souta Wates, Vou. 97, Part 1 A. K. SANGHI AND E. P. BAKER (ab SHEEN, S. J., and Snyper, L. A., 1964.—Studies on the inheritance of resistance to six stem rust cultures using chromosome substitution lines of a Marquis wheat selection. Can. J. Genet. Cytol., 6: 74-82. , 1965.—Studies on the inheritance of resistance to six stem rust cultures using chromo- some substitution lines of Kenya wheat. Can. J. Genet. Cytol., 7: 374-387. SraxMan, E. C., Stewart, D. M., and Lorcrerine, W. Q., 1962.—Identification of physiological races of Puccinia graminis tritici. United States Department of Agriculture. Agriculture Research Service. Bull. H 617 (revised 1962), pp. 53. Watson, I. A., and Lure, N. H., 1959.—Somatic hybridization between Puccinia graminis var. tritici and Puccinia graminis var. secalis. Proc. Linn. Soc. N.S.W., 84: 207-208. , 1963.—The classification of Puccinia graminis var. tritici in relation to breeding resistant wheats. Proc. Linn. Soc. N.S.W., 88: 235-258. , 1966.—Sr 15—a new gene for use in the classification of Puccinia graminis var. tritici. EHuphytica, 15: 239-250. Wittiams, N. D., and Goven, F. J., 1965.—Inheritance of stem rust reaction in Khapli emmer eross. Urop Sci., 5: 145-147. WituramMs, N. D., Goueu, F. J.. and Ronpon, M. R., 1966.—Interaction of pathogenicity genes in Puccinia graminis f. sp. tritici and reaction genes in Triticum aestivum ssp. vulgare “Marquis” and “ Reliance’. Crop Sci., 6: 245-248. PROCEEDINGS OF THE LINNEAN SocieTy oF NEw SoutH WaAtss, Vou. 97, Part 1 EVISCERATION AND REGENERATION IN THYONE OKENI (BELL, 1884) D. J. TRACEY* School of Biological Sciences, University of Sydney, Sydney, N.S.W., Australia [Accepted for publication 17th November 1971] Synopsis The normal anatomy and histology of Thyone okeni was examined in order to compare normal and regenerated tissue. The anatomy was typical of the Dendrochirota, and not significantly different from that of Thyone briareus. The process of auto-evisceration is described, and some explanations of the functional significance of the process are discussed. A new explanation based on nitrogen excretion is considered. Regeneration of new viscera is described, and the mode of regeneration compared with that in other holothurians. The regenerated gut was found to be formed by a proliferation along the whole free edge of the mesentery. The time taken for complete regeneration is greater than six weeks and about four times as slow as in Thyone briareus. A model of the process of regeneration is proposed, in the context of recent ideas on development. INTRODUCTION Some holothurians will eviscerate themselves in response to noxious stimuli or change of season. Regeneration of the viscera following auto-evisceration has been studied in the aspidochirotes Stichopus (Bertolini, 1930 ; Dawbin, 1949) and Holothuria (Bertolini, 1932; Kille, 1936) and in the dendrochirote Thyone briareus. Several workers have studied evisceration and regeneration in this species (Pearse, 1909; Scott, 1914), and Kille (1935) gave a comprehensive picture of its morphological changes during regeneration. In the aspidochirotes, even closely related genera show quite different modes of regeneration and so in the work to be described, the process of regeneration was followed in a species of dendrochirote not previously examined, in order to establish whether it followed a course of regeneration similar to that described for Thyone briareus by Kille (1935). MATERIALS AND METHODS MATERIAL Occurrence. Thyone okeni (Bell, 1884) occurs in about 16 feet of water off Bottle and Glass rocks, Port Jackson, N.S.W. Each animal is invested with a coat of large pieces of shell and gravel and clings to the underside of mussel clumps with its tube feet. Classification. This animal was classified according to the artificial key given by Clark (1946). Spicules were prepared by macerating the whole body wall in hot 10°% KOH, and were then dehydrated and mounted. They are com- pletely unornamented spines (Fig. 1 (B)), few in number and quite atypical for holothurians in general, and indicate Thyone okeni Bell—of which only three Specimens have been previously reported (Bell, 1884; Clark, 1921). In fact Clark found no spicules apart from the tables in the tube feet (Fig. 1 (A)). * Present address: Department of Biological Sciences, Stanford University, Stanford, California 94305, U.S.A. PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WatsEs, Vou. 97, Part 1 Orel mm 74 EVISCERATION AND REGENERATION IN THYONE OKENI EXPERIMENTAL PROCEDURES Some 30 animals were collected and kept in large aquaria, in which the water was kept circulating for aeration at the surface. The sea water was filtered and. -changed every two days for the duration of the experiment. The water temper- ature was constant at 19-++1° C. Two methods of evisceration were tried—ammoniated sea water (Kille, 1935) and injection of distilled water (Dawbin, 1949). For the study on regeneration, 16 animals were induced to eviscerate by injection of about 0-3 ml. distilled water per gramme wet weight. Wet weight is quite variable in each animal, but the amount of water used is not critical. In every case the lantern, gut, and as much gonad as possible was eviscerated ; this was ensured by washing the animals through with extra water. The animals were kept for periods of up to eight weeks, during which six (37°) died and were discarded. Histolysis and decomposition is very rapid after death. Every week one or two of the animals were narcotized by gradual addition of MgSO, to 100ml. of surrounding sea water. They were then killed by dissection and immersion in Bouin’s fixative. Specimens were left for a week in Bouin’s for adequate fixation and some decalcification. Every animal was displayed, examined and drawn under the dissecting microscope, and three transverse slives of tissue taken for histological examina- tion. One was taken in the area of the developing lantern, one in the region of the small intestine, and one showing the large intestine. About 100 serial sections were taken from the first slice, while 40 sections were taken at selected intervals through the second and third slices. Sections were 8y thick and ordering was retained throughout. All sections were stained with Lillie’s modification of the“triple Masson stain. RESULTS NorMAL ANATOMY (Fig. 2) The normal anatomy of Thyone okeni differs little from that of Thyone briareus, but the features important in regeneration will be outlined. The animal is brown and about 50 mm. long, and tube feet are scattered profusely and uniformly over the whole body, so that the ambulacra can hardly he distinguished. The aquapharyngeal bulb is about 20 mm. long, and is supported by retractors whose insertion is nearly as far back as the middle of the body. The genital tubules are numerous, slender and long, and inserted behind the middle of the body. There is a single polian vesicle and madreporite. The course of the gut is typical for holothurians. From the pharynx, the descending small intestine leads down the edge of the dorsal mesentery to the posterior of the body cavity, where it recurves forwards and becomes the ascending small intestine, supported by the left mesentery. Before it reaches the anterior of the animal, it turns once again to become the large intestine, supported by the ventral mesentery and leading into the cloaca. There are altogether six regions of the alimentary canal, distinguishable morphologically and histologically (Nace, 1966). These are the pharynx (inside the lantern), then a short oesophagus, a muscular stomach, a long and coiled small intestine forming the bulk of the gut, the rectum which is a dark brown expansion in contrast to the prevailing light orange, and finally the pulsatile cloaca. The relative lengths of these regions ean be seen in Fig. 2. Transverse sections of the gut show a lining epithelium, an inner connective tissue layer, a muscle layer, an outer connective tissue layer and coelomic epithelium. The layers differ in their proportions and somewhat in their staining along the gut, and there is a cuticle lining the stomach. The rectum carries large numbers of brown globules in the inner connective tissue layer and lining epithelium. PROCEEDINGS OF THE LINNEAN SocrETY or NEw SoutH Wates, Vou. 97, Part 1 D. J. TRACHY 16, CMV MC | RC Fig. 2. Thyone okenti—normal anatomy. OH oesophagus, RM=retractor muscle, MP=madreporite, LN=lantern, PV=polian vesicle, GD—=—gonoduct, RT=respiratory tree, ST=stomach, SI=—small intestine, CMV=contra mesenterial cross blood vessel, MC=cord of mesentery supporting left respiratory tree, LI=—large intestine, IMV=ipsi mesenterial cross blood vessel, RC=rectum. PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SoutH Watss, Vou. 97, Part 1 76 EVISCERATION AND REGENERATION IN THYONE OKENTI The cloaca bears two respiratory trees which continue to function after evisceration. The only other significant structure in the body cavity is the gonad tuft—this is a large mass, purple in the female and light brown in the male, which may fill half the body cavity. HVISCERATION Kille’s ammonia treatment did not succeed in inducing evisceration. Injection with distilled water was effective in every case, although there was wide variation in sensitivity. The time lapse between injection of water into the body cavity and the start of evisceration varied between 0 and 20 min. with an average of about 5 min. After this time, contraction of the body wall raises the coelomic fluid pressure until the tentacles are everted. Further contraction distends the proximal thin-walled part of the introvert. This bursts and coelomic fluid rushes out carrying with it the base of the lantern. Meanwhile the retractor muscles have broken, and the tentacle crown becomes further separated from the rest of the body. The gut is extruded, still enclosing some mesentery and the hemal plexus in its loops. Lastly, variable amounts of gonad are expelled, and the lantern and tentacular crown become completely detached. Eventually the circular muscles around the hole contract and close off the body cavity. The animal has now lost all means of feeding or digestion, and the associated blood system, as well as the central nervous and water vascular systems. It is left with the body wall, the cloaca and respiratory trees and most of the mesentery. REGENERATION Lantern. The lantern is a complex structure, and because the series of specimens was limited and rather variable, the process of regeneration could not be recon- structed as completely as Kille did with Thyone briareus. However, general features of the process were the same in both species. At first a connective tissue platelet heals over the site of the wound, and this supports the lantern rudiment, which is a flattened disc penetrated by the edge of the dorsal mesentery. At two weeks the longitudinal water canals have grown forward and unite to form a water vascular ring with a diameter of about 1mm. The lantern is still wider than long, but by four weeks it has extended and differentiated until it is 3-5 mm. long and 1mm. wide. At six weeks (Fig. 3 (D)), the Jantern is 8mm. long and 3mm. wide and differentiation is complete, even to the polian vesicle and the madreporite with its stone canal. Meanwhile, the longitudinal muscles have grown forward with the underlying water canals towards the introvert. At this point the inner part of the longi- tudinal muscle constricts and eventually pinches off to form a separate strand. This strand is still connected with the longitudinal muscle by a temporary mesentery, continuous with connective tissue which can be seen in sections to extend between the inside and outside of each longitudinal muscle. As time goes on, the new retractor muscle extends its origin posteriorly and the supporting mesentery disappears, until at six weeks the retractor muscles are distinguishable from the normal only in their slenderness and more posterior/anterior origin. Regeneration of Gut. The two main features of gut regeneration are the develop- ment of a new gut from the proliferating mesentery edge, and changes in the distribution of the mesentery which at first reduce the length of the free edge and then gradually return it to its original length. Initially the width of the mesentery in the posterior elbow is about 4 mm. and the length of the free edge is 145mm. By two weeks the mesentery width has increased greatly in the posterior (15 mm.) and anterior elbows in such a way PROCEEDINGS OF THE LINNEAN SocieTy oF New SoutH Wates, Vou. 97, Part 1 Fig. 3. (B) 14 days. (C) 21 days. (4) 42 days after evisceration. D. J. TRACEY U0 Ee Onin A B C D Thyone okeni—stages in the regeneration of the gut from mesentery edge. (A) 7 days. PROCEEDINGS OF THE LINNEAN Socrety or NEw SoutH Waltzes, Vou. 97, Part 1 78 EVISCERATION AND REGENERATION IN THYCNE OKENI that the free edge of the mesentery has been reduced in length (50 mm.). The course of the gut is now virtually straight from the anterior connective tissue platelet to the intact cloaca. At this stage, the new gut is visible under the dissecting microscope as a distinct thickening (¢. 0:-1mm.). This thickening consists of a bead of connective tissue enclosed by epithelium continuous with that of the mesentery (Fig. 4 (A)). The gut rudiment varies in thickness along the Gsilas O