i" pense aag nn ere te Trey RN tAlNee Te NOU leh gin beer eaewonete renew rity ~ vere cy pete a gia eber’ aaewomete Tense TTY nate o Mngeeyriewrarmeasnre yee ay es aearied aya ee oe appa gveneeete tree ete wrnee™ er = eo nt re tarts sei ~~ Syren soe ann Poem b oA PPR OT ee Oo adios obiackins ore oe ener ee wpe: a senaaoet aiakee qo treme ae eee ee Pare 2S Og te PTO “NA ~- “ SS tare et = a tee My in Heol ¥ t OE aby oe bs ify, Meaty Proceedings of the Linnean Society of New South Wales VOEUME 9 Nos. 410-412 CONTENTS OF PROCEEDINGS, VOLUME 91 PART 1 (No. 410) (Issued 5rd November, 1966) (Presidential Address and Papers read March—April, 1966) CONTENTS Page Annual General Meeting Report on the Affairs of the Society for the Year .. 1 Elections We ae ave : 3 Obituary notices 4 Balance sheets it D. T. ANDERSON. Presidential Address. The comparative early embryology of the Oligochaeta, Hirudinea and Onychophora ll) R. BASDEN. The composition, occurrence and origin of lerp, the sugary secretion of Hurymela distincta (Signoret) ha ba $4 we AR: A. A. MARTIN and M. J. LITTLEJOHN. The breeding biology and larval development of Hyla jervisiensis (Anura: Hylidae). (Plate I) .. 47 R. C. CARoLIn. Seeds and fruit of the Goodeniaceae .. is oe D. M. GRIFFIN. Fungi attacking seeds in dry seed-beds Pe «ey 84 R. Wass. On the species Fenestella horologia Bretnall and Minilya duplaris Crockford. (Plate IT) a = a Re a a so MD) R. Wass. Two new species of Permian brachiopods from Queensland. (Plate ITI) .. a ae ae ee ae zs Be eG PART 2 (No. 411) (Issued 22nd December, 1966) (Papers read June-July, 1966) CONTENTS D. G. CATCHESIDE. Sir William Macleay Memorial Lecture, 1966. The Centenary of Mendel : , : AotA M. RicHAarps. The Rhaphidophoridae (Orthoptera) of Australia. 4. A new genus from South Australia G. M. PuHuip. Notes on three recently ee Australian Tertiary echinoid genera a 28 f ie ay ¥ A. T. Hotcuxiss. Chromosome numbers in Lamprothamnium .. N. V. DoBROTWoORSKY. Mosquitoes of Tasmania and Bass Strait Islands. (Plates IV-V) Mary D. TINDALE. New taxa of Acaeia from Eastern Australia. No. 2 EK. B. Britton. (Communicated by Dr. D. F. Waterhouse.) The applica- tion of the names Sericesthis pruinosa (Dalman) and Sericesthis nigrolineata Boisduval (Coleoptera : Scarabaeidae: Melolonthinae). . Page 101 109 114 118 121 147 152 PART 3 (No. 412) (Issued 22nd March, 1967) (Papers read September—November, 1966) CONTENTS EK. P. BAKER and Y. M. UPADHYAYA. Studies on the inheritance of rust resistance in oats. IV. Expression of allelomorphism and gene interaction for crown rust resistance in crosses involving the oat variety Bond E. F. Rix. A new corallanid isopod parasitic on Australian freshwater prawns B. P. Moort. New Australian cave Carabidae (Coleoptera) PATRICIA Kort (Mrs. W. B. MATHER). Atopozoa deerata (Sluiter): a discussion of the relationships of the genus and species EH. P. BAkER. Inheritance of resistance to bunt in Turkey wheat selections R. DomrRow. RKhinonyssine nasal mite infestations in birds at Mitchell River Mission during the wet and dry seasons HELEN P. RAmsay. Intraspecific polyploidy in ee ay rotulatum (Hedw.) Brid. (Plates VI-IX) is : : B. RK. Hewitt. Highlands peat of the Malayan Peninsula Abstract of Proceedings List of Members List of Plates List of New Genus, New Species and New Subspecies Index CORRIGENDUM Page 155 176 179 185 189 211 220 231 233 241 249 249 250 Page 110, line 37.—For Vovotettix naracoortensis, nu.sp. read Novotettiix naracoortensis, D.Sp. SYDNEY PRINTED AND PUBLISHED FOR THE SOCIETY BY AUSTRALASIAN MEDICAL PUBLISHING CO. LTD. Seamer Street, Glebe, Sydney and. SOLD BY THE SOCIETY 1967 ANNUAL GENERAL MEETING 30TH MARCH, 1966 The Ninety-first Annual General Meeting was held in the Society’s Rooms, Science House, 157 Gloucester Street, Sydney, on Wednesday, 30th March, 1966, at 7.30 p.m. Dr. D. T. Anderson, President, occupied the chair. The minutes of the Ninetieth Annual General Meeting (31st March, 1965) were read and confirmed. REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR The Society’s Proceedings for 1965, Vol. 90, Parts 1 and 2, were published on Sth November, 1965, and 18th February, 1966, respectively. Part 3 has yet t- be issued. The format of the Proceedings has been changed with Vol. 90 end there appears to be general agreement that the new, more modern style is a great improvement. All concerned with the alterations are to be con- eratulated on the result. The subscription price of the Proceedings is now $11.40 per annum, including postage. A revision of prices for available Volumes and Parts of the Proceedings, postage included, has been made as follows :— 1st Series (Vols. 1-10) (1875/6—1885), $416.40 per Volume; 2nd Series, Vol. 1 (Vol. 11) (1886) onwards, $411.40 per Volume; each Part, $A5.15; $A4.15 each for current numbers, and, for Volumes of five or six Parts, $A1.15 each for the first and last Parts. During the year 16 new members were admitted to the Society, three died, seven resigned and two were removed from the list of members. The numerical strength of the Society at 1st March, 1966, was: Ordinary Members, 270; Life Members, 32; Corresponding Member, 1; total, 303. It is with regret that the deaths: of the following members are recorded : Mr. G. H. H. Hardy, on 9th January, 1966; Miss Florence Sulman, M.B.E., on 15th June, 1965; and Dr. A. R. Woodhill, on 27th July, 1965. (See page 4 for obituary notices.) Papers read at Ordinary Monthly Meetings totalled 26. Lecturettes were given at the following meetings: April, Some Aspects of Coastal Morphology in New South Wales, by Mr. J. R. Hails; June, Trace Fossils: their Classification and Palaeoecological Significance, by Dr. B. D. Webby; July, Ecology of Ruppia and Zostera, by Mr. R. Higginson; September, Opossums, by Dr. M. P. Marsh, and October, Some Problems of Evolutionary Theory, by Dr. Paul Ehrlich. Interesting discussions followed the lecturettes. We are grateful to these lecturers, who contributed greatly to the interest of the meetings. At other meetings notes and exhibits were presented. No meetings were held in May or August. Dr. N. G. Stephenson, Mr. L. A. 8S. Johnson and Dr. Erik Shipp were elected members of Council in place of Professor B. J. Ralph, Professor W. L. Waterhouse and Professor I. A. Watson, who had resigned. Two Special General Meetings were held to consider the recommendations from the Council that Rules V and VI be altered to read as follows : Rule V.—No person declared elected under Rule IV shall be admitted to any of the privileges of membership until he has signed a written acceptance of membership and paid his first Annual Subscription. Rule VI.—The Annual Subscription shall be three pounds ten shillings (seven dollars) and shall become due in advance on the first day of March in each year, or, in the case of New Members, immediately on election ; provided that Ordinary Members elected after the first day of October in any year shall have the option of paying the Annual Subscription either for that year or in advance for the following year. Any Ordinary Member who has paid the Annual Subscription for forty years shall be exempt from payment of further subscriptions. A 2 PRESIDENTIAL ADDRESS The recommendations were unanimously adopted on 29th September and confirmed on 27th October, 1965. Library accessions from scientific institutions and societies on the exchange list amounted to 2,016 compared with 2,222 and 2,174 for the years 1964 and 1963 respectively. The total number of borrowings of books and periodicals from the library by members and institutions for the year was 253. Members and others continued to consult publications in the Society’s rooms, and books and periodicals were made available for photographic copying. The following requests for exchange of publications were acceded to during the year: One copy of the Proceedings to the Institute of Botany and one to the Institute of Zoology, Academia Sinica, Nankang, Taipei, Taiwan, Republic of China, in return for their Botanical Bulletin and Zoological Bulletin respectively ; Proceedings to the Central Library of the Academy of Science of Kazakh, Alma Ata, Kazakh, SSR., for the publications of the Botanical and Zoglogical Institutes ; Reprints of geological papers to the All Union Geological lL brary, Leningrad, U.S.S.R.; Botanical reprints to the Academie des Sciences de Bulgarie, Sofia, Bulgaria, in return for its Bulletin of the Institute of Botany ; and Botanical reprints, as a gift, to Estudos Gerais Universitarios de Mocambique, Laboratorio de Botanica, Lourenco Marques, Mocambique, as from 1965. The recently-formed Australian Society of Limnology sends its Newsletter to the Society’s library. The Society now sends its Abstract of Proceedings and reprints of any papers required, instead of the Proceedings, to the Nasionale Museum, Bloemfontein, South Africa. The exchange of publications with the Universidad de Buenos Aires, Buenos Aires, Argentina, has been cancelled. The Lloyd Library and Museum, Cincinnati, U.S.A., was removed from the Society’s exchange list owing to the publication ‘‘ Lloydia”’ being no longer available by exchange as from 1966. The total revenue accruing to the Society from its one-third ownership of Science House was $2,244.18 for the year ending 31st August, 1965. On 8th September, 1965, a lecture on Radio-carbon Dating in the Quaternary was delivered by Dr. Harry Godwin, Professor of Botany, University of Cambridge, under the joint auspices of this Society and the School of Biological Sciences, University of Sydney. During the year the Society received as a gift from the Linnean Society of London a portrait of Linnaeus. This painting, copied from the Roslin portrait presented to the Linnean Society of London in 1814 by Joseph Sabine, one of the original Fellows of the Society, hung for many years in the meeting- room of the Society behind the President’s chair. Further particulars of this portrait may be found in Proceedings Linnean Society, London, Session 118, 1905-1906: 68. A conversazione, kindly arranged by Mr. Ronald Strahan, was held on the afternoon of Saturday, 18th September, 1965, in the Zoology Department, University of New South Wales, when material and apparatus were exhibited to an appreciative gathering by members of the Society and others engaged in scientific research. A Conservation Photographic Exhibition arranged by the Society was put on show at the Australian Museum with the kind co-operation of the Director on 24th March, 1965. The photographs were subsequently shown in Tasmania and elsewhere and eventually presented to the Fauna Protection Panel. The Council supported the nominations of Dr. R. C. Carolin and Professor J. LeGay Brereton of the University of New England, to fill two vacancies on the Fauna Protection Panel and these two gentlemen were appointed by the Chief Secretary. During the year Council, finding an increasing number of matters involving Nature Conservation coming before it, appointed a Conservation Committee to consider and make recommendations concerning such matters. Already Council has made representations to the appropriate State Ministers regarding the PRESIDENTIAL ADDRESS 3 biological resources of the Clarence River valley in relation to flood-mitigation and water-conservation plans, the desirability of a Myall Lakes Reserve, and the appointment of an ecologist to fill a vacancy on the Kosciusko State Park Trust. Innnean Macleay Fellowship During the year Mr. A. J. T. Wright, B.Sc., the Linnean Macleay Fellow in Palaeontology, studied the Devonian sediments and faunas of the Mudgee district and the Capertee Valley area. A paper entitled ‘‘ Cerioid Stringo- phyllidae (Tetracoralla) from Devonian strata in the Mudgee district, New South Wales ” will appear in the Society’s Proceedings, Vol. 90, 1965, Part 3. Owing to his appointment as Lecturer in Geology in Victoria University, Wellington, New Zealand, Mr. Wright did not seek reappointment. No appointment for 1966 was made. Linnean Macleay Lectureship in Microbiology Dr. Y. T. Tchan, Reader in Agricultural Microbiology and Linnean Macleay Lecturer in Microbiology, University of Sydney, reported on his work for the year ending 3ist December, 1965, as follows: Progress has been made on the influence of higher concentration of acetate on the calcium requirement: of Azotobacter. In collaboration with the Biochemical Department of the University of New South Wales the cytochemical studies of Azotobacter have been started, using strictly controlled cultural conditions. Advances have been made on the serological reaction of Azotobacter. It has been shown that antigenic properties of Azotobacter vary with ecological environment of the culture. On the study of herbicides, a new quick assay method has been designed and could be useful for field trials. The Honorary Treasurer (Dr. A. B. Walkom) presented the balance sheets for the year ending 28th February, 1966, duly signed by the Auditor, Mr. 8. J. Rayment, F.C.A., and his motion that they be received and adopted was carried. unanimously. PRESIDENTIAL ADDRESS The Comparative Early Embryology of the Oligochaeta, Hirudinea and Onychophora On morphological and embryological grounds, the Onychophora (Peripatus) can be presumed to have evolved from a group of soft-bodied, metamerically segmented coelomates, usually referred to as lobopods. In terms of adult morphology, existing annelids do not fit the requirements of the hypothetical lobopod, and the possibility of including the ancestors of the Onychophora within the annelids has generally been rejected. A new comparison of onychophoran embryonic development with that of oligochaetes and leeches, however, reveals that onychophoran development can be interpreted as annelid development in a modified form. The extinct lobopods must therefore have had annelid-like embryos, which make it probable that they were paraphyletic with annelids. (For full text see pp. 10 et seq.) No nominations of other candidates having been received, the President declared the following elections for the ensuing year to be duly made: President: R. C. Carolin, Ph.D., B.Sc., A.R.CS. Members of Council: D. T. Anderson, B.Se., Ph.D.; RB. C. Carolin, Ph.D., B.Sc., A.R.C.S.; L. A. S. Johnson, B.Sc.; E. Shipp, Ph.D.; N. G. Stephenson, M.Se., Ph.D.; and A. B. Walkom, D.Sc. Auditor: S. J. Rayment, F.C.A. The President then installed Dr. R. C. Carolin as President. A cordial vote of thanks to the retiring President was carried by acclamation. OBITUARY NOTICES GEORGE HUDDLESTON HURLSTONE HARDY Mr. Hardy, who died on 9th January, 1966, had been a member of the Society since 1917. He was born in Twickenham, London on August 14th, 1882. He received an early training in the engineering profession, but soon developed an interest in entomology which remained with him all his life. He came to Perth, Western Australia, in 1911, remaining there for two years,and was then appointed to a position with the Tasmanian Museum, Hobart, which he held for five years. He arrived in Sydney in 1918 where he spent the following four years,»and in 1922 was appointed Walter and Eliza Hall Fellow in Economic Biology at the University of Queensland. This position was held until 1932 and in 1949 he left Brisbane taking up residence at Katoomba, N.S.W. Due largely to his efforts the Entomological Society of Queensland was formed in 1924. He became its first Treasurer, holding this position for many years. Much of his scientific life time was devoted to the study of Diptera, and in all he published 144 papers on the biology and systematics of Australian Diptera. He contributed 40 papers to the Society’s Proceedings between 1920 and 1959 and one (with T. H. Johnston) in 1923 entitled ‘‘ A revision of the Australian Diptera belonging to the genus Sarcophaga”’. He also published papers in Papers and Proceedings of the Royal Society of Tasmania, Proceedings of the Royal Society of Queens- land, Records of the Australian Museum, Australian Zoologist and Queensland Agricultural Journal, and in overseas journals including the Annals and Magazine of Natural History, Bulletin of Entomological Research, Stylops, Nature, Entomologist’s Monthly Magazine and Proceedings of the Royal Entomological Society of London. Mr. Hardy took a keen interest in the Society during his stay in Sydney and Katoomba and attended meetings when possible. The sympathy of members of the Society is extended to his widow and married daughter. The following papers were published by him in addition to those given in the Bibliography of Australian Entomology, 1775-1930, by Anthony Musgrave :— 1931. Aphididae in Australia. Proc. roy. Soc. Q’land, 43: 31-36. On genus Damaromyia Kertesz. Ann. Mag. nat. Hist., (10), 8: 120-128. 1932. Two new Australian species of Pollenia. Proc. Linn. Soc. N.S.W., 57: 338-340. Some new Australian Sarcophagid flies and notes on others. Aust. Zool., 7: 275-281. Australian flies of genus Actina (Stratiomyiidae). Proc. roy. Soc. Q’land, 43: 50-55. Notes on Australian Stratiomyiidae. ibid. 44: 41-49. : Notes on Sarcophaga peregrina-group. Bull. ent. Res., 23: 45-48. Some Australian species of Calliphora (subgenera Neopollenia and Proekon). ibid. 23: 549-558. 1933. Miscellaneous notes on Australian Diptera. I. Proc. Linn. Soc. N.S.W., 58: 408-420. Notes on Australian Syrphinae. Proc. roy. Soc. Q’land, 45: 12-18. 1934. Miscellaneous notes on Australian Diptera. II. Empididae. Proc. Lin. Soc. N.S.W., 59: 173-178. Notes on Sarcophagid flies. Aust. Zool., 8: 50-53. Notes on Australian Muscoidea. Proc. roy. Soc. Q’land, 45: 30-37. The Asilidae of Australia. Part 1. Ann. Mag. nat. Hist. (10), 13: 498-525. The Asilidae of Australia. Part 2. ibid. (10), 14: 1-35. The genus Tabanus in Tasmania. Stylops, 3: 43-48. 1935. Miscellaneous notes on Australian Diptera. III. Chrysosomatinae. Proc. Linn. Soc. N.S.W., 60: 248-256. The Asilidae of Australia. Part 3. Ann. Mag. nat. Hist. (10), 16: 161-187. The Asilidae of Australia. Part 4. esuodsng 69°EL2‘6 ee vie oe qunoodoy euLooUuTyT ean mebaaleay fae 99°FZ8'T st ++ gumnosay Surpurqyoog. 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ANDERSON School of Biological Sciences, University of Sydney [Delivered 30th March, 1966] Synopsis The primitive, large, yolky eggs of tubificid and lumbriculid oligochaetes and glossiphoniid leeches undergo a modified spiral cleavage yielding a yolky blastula. The pattern of presumptive areas in the blastula wall is interpretable as a modification of the polychaete presumptive area pattern, in association with increased yolk. The same pattern of presumptive areas is retained in the secondarily microlecithal embryos of lumbricid oligochaetes, gnathobdellid and pharyngob- dellid leeches and probably other clitellates, in spite of further cleavage modifications and of embryonic adaptations to albumenotrophy. The primitive, large, yolky eggs of ovoviviparous Onychophora undergo centrolecithal cleavage, setting out a blastoderm around a central yolk mass. In viviparous non-placental species, although yolk is absent and cleavage is modified, a similar blastoderm is usually formed around a central cavity. In viviparous placental species, cleavage is highly specialized, yielding a small blastodermic vesicle at the end of a placental stalk and plate. The blastodermal presumptive areas of Onychophora, like those of clitellates, retain a constant pattern in spite of secondary yolk loss and its attendant modifications. Taking into account the events of gastrulation, the blastodermal presumptive area pattern of Onychophora is interpretable as a modification of the primitive clitellate presumptive area pattern, in association with a further increase in yolk. It is suggested that a clitellate-like mode of early embryonic development must have characterized the ancestors of the Onychophora, and that these ancestors were members of the spiral cleavage assemblage and were probably paraphyletic with the ancestors of annelids. INTRODUCTION Following a review of the embryology of polychaete annelids by Anderson (1966a), the embryonic development of clitellate annelids and the modifications which they show as compared with polychaetes now become a subject of renewed interest. While the development of yolky polychaete eggs always retains traces of a primitive mode of development from a small, microlecithal egg through a planktotrophic trochophore and metamorphosis, that of clitellates shows no such traces. As is well known, both oligochaetes and hirudineans deposit their eggs in a cocoon in which development proceeds directly to hatching as a miniature adult with numerous segments. Primitively, the embryo develops from a relatively large yolky egg, the cleavage, gastrulation and organogeny of which are adapted to the presence of voluminous yolk reserves. Secondarily, in the majority of clitellate families, the egg is small and microlecithal and the embryo is adapted to feeding on the albuminous contents of the cocoon. The nature of both types of modification has hitherto yielded no clue as to the possible marine antecedents of primitive clitellate development or the derivation of the secondary yolkless from the primitive yolky condition. When the data of clitellate embryology are reexamined in the light of new understanding of polychaete development, however, solutions to both of these problems can be proposed, as will be shown below. Primitive clitellate development exemplifies a particular modification of spiral cleavage development to voluminous yolk and the production of a metamerically segmented annelid. The embryonic development of Onychophora is also primitively adapted to voluminous yolk, and to the formation of a metamerically segmented animal with a number of annelid-like features. In the absence of ‘ spiral cleavage’ features in onychophoran development, the PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SoutH WALES, Vol. 91, Part 1 D. T. ANDERSON ial derivation of Onychophora from a spiral cleavage ancestry has hitherto proved elusive of recognition (Manton, 1965) but, as will also be shown below, pursuance of the comparison in an appropriate way reveals the development of the Onychophora, not only as a modification of spiral cleavage development, but also as one with unmistakable clitellate-like antecedents. It is true that certain difficulties attend this line of argument. The descriptive data on the embryology of the Oligochaeta, Hirudinea and Onychophora are incomplete in many critical ways and few of the available papers embody results obtained by modern histological techniques. Among the oligochaetes for example, T'ubifex rivulorum and Rhynchelmis limosella are the only species with large, yolky eggs whose development has been adequately described (Schmidt, 1922; Penners, 1922, 1924; Swetloff, 1923b; Meyer, 1929), all other investigations being concerned with the specialized embryos of naidids, Bdellodrilus and earthworms. The same situation obtains for the Hirudinea, where again only two forms with large yolky eggs, Glossiphonia and Theromyzon, have been investigated (Schliep, 1914; Schmidt, 1917, 1925a), the remaining descriptions being confined to specialized piscicolid, gnathobdellid and pharyngobdellid development. The Onychophora present a further difficulty in that nothing is known of the development of the large, yolky eggs of oviparous onychophorans and only a little of the yolky eggs of ovoviviparous species (Sheldon, 1888, 1889a; Evans, 1902). On the other hand, the specialized development of the viviparous Onychophora, especially as described for Peri- patopsis by Manton (1949), provides many inferences which are of value for comparative purposes, and permits a picture to be drawn of clitellate and onychophoran development which holds much of interest for the phylogeneticist and epigeneticist. CLEAVAGE Primitive cleavage in the Oligochaeta In Tubifex and Rhynchelmis, the spiral pattern of cleavage is retained, but is modified in ways related to the magnitude of the large, yolky egg and its development in a protective cocoon. The nature of these modifications has hitherto been obscured by employment of the Wilsonian system of enumeration of spiral cleavage (Wilson, 1892), which places emphasis on the integrity of quartettes and individuality of single blastomeres and their fate. If, as for polychaetes (Anderson, 1966a), attention is alternatively directed towards the formation and subsequent fate of presumptive areas in the wall of the blastula, the oligochaete response to yolk is much more clearly discerned. Both in Tubifer (Penners, 1922, 1924) and Rhynchelmis (Vedjovsky, 1886, 1888-92 ; Bergh, 1890; Penners, 1922; Swetloff, 1923), the first two cleavage divisions are at right angles along the animal-vegetal axis of the egg, the next four perpendicular to this axis and, except for a precocious onset of bilateral cleavage in the D-quadrant, successively dexiotropic, laeotropic, dexiotropic and laeotropic in the classical spiral cleavage sequence. In association with the large size of the egg, 300—500u in diameter in Tubifex, 1000u in Rhynchelmis, the first division is very unequal, AB being much smaller than CD, and the second division maintains this inequality in the D-quadrant, D remaining large, while C is only a little larger than A and B (Figs 1, 2). In polychaetes with yolky eggs, where the D cell is also disproportionately large, much of it is subsequently segregated into a ring of large ectoteloblasts and a pair of large M-cells within the ring, forming a growth zone which gives rise to several trunk segments before the yolk reserves of the embryo are exhausted (Anderson, 1966a). The large D cell in primitive oligochaetes is a developmental adaptation subserving a similar function. In Tubifex, for example, 34-38 segments are formed before the embryo hatches from the cocoon. The orientation of the four quadrants in Tubifex at the 4-cell stage is as in polychaetes, D being dorsal, and A, B and C left-lateral, ventral and right-lateral respectively relative to the anteroposterior axis of the embryo, though there 12 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA is a change in quadrant orientation as development proceeds further. In Rhynchelmis at the 4-cell stage, on the other hand, the large D cell is posterior in position, with A on the left, B anterior and C on theright. Cleavage in Tubifex finally results in displacement of B-quadrant cells to an anterior and D-quadrant cells to a posterior position as described below. In Rhynchelmis quadrant reorientation, presumably through a redistribution of ooplasms, manifests itself in the egg. Associated with the large size of the D cell, early cleavage after the 4-cell stage in Tubifer and Rhynchelmis differs from that of polychaetes in a marked tendency to precocious division of the D-quadrant cells, and to a lesser extent of the C-quadrant cells, so that the cells of each quartette cut off from the stem cells at the 3rd and subsequent divisions are formed in succession, the D-quadrant leading, the A and B quadrants lagging. Figs 1-4.—1, Tubifex, 4-cell stage, anterior view with dorsal surface on the right. 2, Rhynchelmis. 4-cell stage, dorsal view with posterior end on the right. 3, Tubifex, 8-cell stage, anterior view, 4, Rhynchelmis, 8-cell stage, dorsal view. (After Penners, 1922; Swetloff, 19236.) The key to the shading conventions employed in Figs 1-33 is provided by Fig. 13. At the 3rd cleavage division (Figs 3, 4), dexiotropic and very unequal, a ist quartette of very small cells la—ld is cut off from stem cells 1A-1D. In Tubifex, 1a-1d lie bilaterally on either side at the anterior pole, 1d and la to the left, 1b and 1c to the right. In Rhynchelmis, they are similarly bilateral, but dorsally placed. The laeotropic 4th division begins with the division of 1D into 2d and 2D (Fig. 6). 2d is much larger than any cell of the 1st quartette, though in Tubifex it is only about one-quarter, and in Rhynchelmis less than one-quarter, of the size of 2D. In Tubifex, 2d pushes forward anterodorsally between 1d and le to lie in the dorsal midline, with the 1st quartette cells as an are around its anterior face. In Rhynchelmis, 2d lies mid-dorsally behind the 1st quartette. 2a, 2b and 2c are now cut off laeotropically as small cells, so that 2a is added to the left, 2b to the front and 2c to the right of the micromere are bordering 2d (Figs 5, 6). D. T. ANDERSON 13 At the same time, the 5th division begins in the D-quadrant. The products of this division as it affects 2D differ in the two embryos. In Tubifex, the d component of the 3rd quartette, 3d, is a small cell cut off dexiotropically and added to the left-hand end of the micromere are (Fig. 5). In Rhynchelmis, 3d is cut off in the dorsal midline as a cell directly behind and equal in size to 2d (Fig. 6). The latter cell, on the other hand, behaves similarly in both species at this stage, dividing unequally, with its smaller daughter pushing into the micromere are while its larger daughter remains mid-dorsal. This is repeated at least twice in subsequent divisions, a cell being added on each occasion to the micromere are, leaving the large cell in the mid-dorsal line. The final designation of the large cell as 2d!" in Tubifex and 2d" in Rhynchelmis (Figs 7, 8) does not alter the fundamental similarity of the process in the two species. Pe “F aC, | 20° Figs 5-8.—5, Tubifex, 17-cell stage, dorsal view. 6, Rhynchelmis, 12-cell stage, dorsal view. 7, Tubifex, 22-cell stage, anterolateral view. 8, Rhynchelmis, 30-cell stage, dorsal view. (After Penners, 1922; Swetloff, 19230.) Meanwhile, the 5th cleavage division continues in the remaining quadrants, the stem cells 2A, 2B and 2C cutting off the small cells 3a, 3b and 3c to lie on the left, anteriorly and on the right respectively, at the outer rim of the micromere are (Figs 5, 7). In Tubifex, the cells 1c and 1d, on the right and left in this arc, also divide equally, increasing the number of micromeres. In Rhynchelmis, they are said not to divide again, but the evidence for this is not positive. The later part of the 5th division is accompanied by precocious 6th division in the D-quadrant. The stem cell 3D divides into large 4d and 4D cells lying in midline (Figs 7, 8). In Tubifex, 4d is equal in size to 4D, although almost all of the yolk in the parent 3D stem cell is confined to 4D. In association with its relatively large size, 4d occupies the posterior midline, pushing 4D ventrally and 3A, 3B and 3C forwards. 3B thus comes to occupy an anterior position, while the micromere are in front of 2d is lifted into a dorsal position. In 14 BARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA Rhynchelmis, 4d is much smaller than 4D, the two cells being posterodorsal and posteroventral respectively. In the remaining yolky stem cells, 3A, 3B and 3C, further divisions are equal, simply increasing the number of yolky cells; and 4D now divides in the same way. . After the formation of 4d, the embryos of Tubifex and Rhynchelmis are remarkably similar in general construction. Hach has (Figs 7, 8): (i) ventrally, a number of large yolky cells, the stem cells 3A—3C and 4D, forming a relatively ereater proportion of the whole in Rhynchelmis than in Tubifex ; (ii) postero- dorsally, one (Tubifex) or two (Rhynchelmis) large cells in the midline, segregated via 2d in Tubifex and via 2d and 3d in Rhynchelmis, surrounded anteriorly and laterally by an arc of micromeres cut off from the stem cells as 1st, 2nd and 3rd quartette cells by a similar series of divisions in both species ; (ii) posteriorly in the dorsal midline, a large 4d cell, relatively greater in size in Tubifex than in Rhynchelmis. The orientation of these cells relative to the major axes is attained in Tubifex during cleavage. In polychaetes in which the D eell is disproportionately large, there is a tendency for A and C cells behind the prototroch to be pushed ventrally, and also for the posterior D cell, 4D, to be pushed ventrally as the large 2d and 4d cells are formed, but the anterior part of the embryo in front of the prototroch remains unaffected by this (Anderson, 1966a). In Tubifea, these tendencies are exaggerated and in the absence of a prototroch, associated with a large egg in a protective cocoon, they influence even the anterior end. After the formation of the 1st quartette, the axial relations of the eight cells are still as in polychaetes. Formation of a large 2d cell, however, displaces the posterodorsal stem cell 2D posteriorly and ventrally. Formation of a large 4d cell accentuates this, pushing 4D into a ventral position. The B quadrant stem cell is thereby displaced to the anterior end, so that 2b and 3b are cut off upwards at this end, in front of 2d. Similarly, the A and C quadrant stem cells are rotated forwards and upwards, and cut off 2a and 3a, and 2¢ and 3c, upwards on the left and right of 2d. In this way, the B quadrant becomes anterior, the D quadrant dorsal and posterior, and the A and C quadrants left and right vertical respectively. It can be suggested that loss of the prototroch is prerequisite to this new pattern of modified spiral cleavage. While the prototroch persists (Polychaeta), modifications associated with an increased size of 2d and 4d posterior to the prototroch have little influence on cells anterior to it. With loss of the prototroch, a new quadrant configuration, established during cleavage in Tubifex, becomes possible. In Rhynchelmis, in association with a relatively greater yolk mass, the new configuration develops directly. Primitive cleavage in the Hirudinea From a comparative viewpoint, the important feature of cleavage in primitive leeches is its close similarity in every respect to cleavage in primitive oligochaetes. In Glossiphonia (Schliep, 1914) and Theromyzon (Schmidt, 1917, 1925a), as in the oligochaete Rhynchelmis, the quadrant orientation, B anterior, D posterior, is established after the first two cleavage divisions, and subsequent divisions proceed in a similar manner in the two species. In association with the large size of the egg, 500-1000u in diameter in Glossiphonia complanata, 600 in diameter in Theromyzon, the 1st cleavage division is unequal, although less markedly so than in primitive oligochaetes, AB being only a little smaller than CD. At the next division, AB divides equally into anterior B and left-lateral A, while CD divides into right-lateral C, equal in size to A and B, and posterior D, slightly the largest cell (Fig. 9). The parallel with Rhynchelmis is unmistakable and continues into subsequent divisions. The 3rd division is very unequal, a 1st quartette of very small cells being cut off dexiotropically from the stem cells to lie in bilateral arrangement, dorsally D. T. ANDERSON 15 on either side of the midline, la and 1d to the left, 1b and 1c to the right (Fig. 10). As in oligochaetes, this division occurs first in the D quadrant, then in the C quadrant, with A and B lagging. The laeotropic 4th division begins in the D quadrant. 1D divides into a large but yolky-free dorsal cell, 2d, displaced at this stage to the right, and an only slightly larger, but yolky, 2D cell lying posteroventrally and to the left. The three remaining cells of the 2nd quartette are then cut off as micromeres. 2a on the left, 2b anteriorly and 2c on the right, around the 1st quartette, making up a micromere cap (Fig. 11). Ic Figs 9-12.—9, Glossiphonia, 4-cell stage, dorsal view. 10, Glossiphonia, 8-cell stage, dorsal view. -11, Glossiphonia, 16-cell stage, dorsal view. 12, Theromyzon, 19-cell stage, dorsal view. (After Schliep, 1914; Schmidt, 1944.) In the dexiotropic 5th division (Figs 11, 12), 2D, 2C, and 2A and 2B throw off, in sequence, small 3rd quartette cells 3d, 3c, 3a and 3b, which add to the edge of the micromere cap. 2d also divides unequally, throwing off a small cell 2d1 on to the posterior edge of the micromere cap, the sister cell 2d? remaining large, posterodorsal and displaced to the right, with 3D lying posteroventrally below it and slightly displaced to the left. At the same time, the 4th division is completed in the 1st quartette, whose cells divide approximately equally into mid-dorsal 1a!—1d1 and adjacent 1a?—1d? in the centre of the micromere cap. After the formation of 2d? at the 5th division, the embryos of Glossiphonia and Theromyzon have the same general configuration as those of primitive oligochaetes at a slightly later stage, namely (Fig. 12): (i) ventrally, a number of large yolky cells, 3A—3C ; (ii) posterodorsally, a large cell 2d?, slightly displaced to the right, with a cap of micromeres anterior to it, cut off from the stem cells aS Ist, 2nd and 3rd quartette cells by a similar series of division in both species ; (iii) posteroventrally, slightly displaced to the left, a large cell 3D. 16 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA The only difference between this pattern and that of oligochaetes summarized in the same way above is the lack of subdivision of 3D in leeches into posterior 4d and ventral 4D. The reason for this difference will be explained below in terms which minimize its significance. Primitive fate of the cleavage blastomeres in the Clitellata As in polychaetes (Anderson, 1966a), once the cell 4d has been cut off in oligochaetes from its stem cell, early in the 6th cleavage division, it becomes possible to assign all blastomeres to presumptive areas of specific subsequent fate. In primitive leeches, the cleavage blastomeres can similarly be assigned to presumptive areas by the end of the 5th cleavage division. In both groups, the pattern of presumptive areas is identical. Their most conspicuous feature in comparison with polychaetes is simplification, associated with the absence of presumptive larval organs. The areas comprise (Figs 13, 14, 18): (a) a dorsal area of presumptive ectoderm ; (b) at the anterior edge of this area, in the dorsal midline, an area of presumptive stomodaeum ; (c) a posterior area of presumptive mesoderm, incorporating presumptive germ cells; (d) a ventral area of pre- sumptive midgut. In the absence of presumptive prototroch, the separation of presumptive ectoderm into anterior and posterior parts seen in polychaetes is not found in clitellates. Presumptive ectomesoderm is also absent. stomodaeum midgut yolk sac mesoderm 13 14 Figs 13-14._13, presumptive areas of the blastula of Tubifex. 14, presumptive areas of the blastula of Rhynchelmis. Diagrams based on data from authors listed in the text. For oligochaetes, the designation of the above areas is supported by the work of Penners (1922, 1924), Penners and Stablein (1930) and Meyer (1929, 1931) on Tubifer, Penners (1929) on Peloscolex benedini, and Vedjovsky (1888-92), Bergh (1890), Penners (1922), Swetloff (1923b) and Iwanoff (1928) on Rhynchelmis. Peloscolex, as far as it has been studied, develops in the same manner as Tubifex. Differences between Tubifex and Rhynchelmis occur only in the constitution of the presumptive ectoderm. Of the four areas listed above, two in oligochaetes correspond in every respect to the homologous areas in polychaetes. The presumptive mesoderm is segregated posteriorly in the single cell 4d which, before it assumes its definitive mesodermal fate, throws off into the interior a pair of small cells, the primordial germ cells. 4d then divides equally bilaterally into M; and M,, teloblasts which subsequently bud off paired ventrolateral mesodermal bands (Figs 15, 16). The presumptive midgut is segregated in the large, ventral, yolky cells 3A-3C and 4D. There also appears to be a close correspondence between the presumptive stomodaeal area of oligochaetes and that of polychaetes. Especially in Tubifex, and perhaps also D. T. ANDERSON iN7/ in Rhynchelmis, the anterodorsal micromeres which make up this area stem mainly from 2b and 3b, while 3a can also be implicated in Tubifex (Fig. 7). Although the area as a whole is displaced anteriorly, its similarity in relative position and mode of segregation to the stomodaeal presumptive area of polychaetes is conspicuous (compare Anderson, 19664). The presumptive ectoderm of oligochaetes is made up of one or, in Rhynchelmis, two large, central cells surrounded anteriorly and laterally by an are of micromeres (Figs 7, 8). In Tubifex and Peloscolex, the single central cell, the main product of 2d, divides equally bilaterally into a pair of cells, each of which gives rise by further divisions to a transverse row of four ectoteloblasts (Figs 15-17). In RKhynchelmis, the anterior central cell, the main product of 2d, band yolk sac Figs 15-17.—15, Tubifex, dorsal view of stage with paired M-cells. 16, Tubifew, posterior view of stage during ectoteloblast formation. 17, Tubifex, left lateral view of stage after onset of ectoteloblast activity. (After Penners, 1924.) divides into a pair of cells which migrate laterally, while the adjacent, posterior, central cell, 3d, divides into a pair of cells each of which divides again into three cells in a transverse row with the large 2d cell at the end of it. The eventual product in each case, although resulting from a slightly different series of divisions, is a row of four ectoteloblasts on either side of the midline. Each ectoteloblast now begins to bud off a row of small cells forwards (Fig. 17). While the ectoteloblasts are forming, the arc of micromeres multiplies and spreads (Figs 15-17): (i) back mid-dorsally, as a strip separating the two ectoteloblast groups ; (ii) up towards the mid-dorsal line from its ends, behind the ectoteloblast groups. B 18 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA The backgrowth mid-dorsally finally meets the upgrowth from the sides. When this stage is reached, the following sub-areas can be distinguished within the presumptive ectoderm: (1) the ectoteloblasts, incorporating presumptive material of the prostomium and cerebral ganglia and the major part of the ectoderm of the peristomium and trunk segments, and already beginning to bud off this material as ectodermal bands; (2) the presumptive temporary yolk sac ectoderm, mid-dorsally and laterally, later incorporated into the segmental epithelium ; (3) the presumptive pygidial epithelium, incorporating presumptive proctodaeum, posterior to the teloblasts. This sub-area, as noted above, has a paired bilateral origin and is also of temporary yolk sac function at this stage. In comparison with the presumptive ectoderm of the polychaetes (Anderson, 1966a, fig. 22), the presumptive mouth region ectoderm and telotroch are absent, the presumptive anterior (prostomial) ectoderm is incorporated into the ecto- teloblasts and all extrateloblastic cells have a new presumptive fate as temporary yolk sac cells, although retaining a capacity for forming definitive trunk and pygidial epithelium. The adaptation of the majority of the ectoderm to a temporary yolk sae function is associated with the large volume of presumptive midgut soon to be accommodated within the interior of the embryo. Deletion of the telotroch, as of the prototroch, requires no explanation. Deletion of distinct mouth region ectoderm and incorporation of presumptive prostomium into the ectoteloblasts can be regarded as a simplification made possible by advanced lecithotrophy, based on the ectoteloblast pattern of polychaetes. The presumptive ectodermal pattern of oligochaetes is thus derivable from that of polychaetes, completing the correspondence of all presumptive areas and at the same time revealing a new presumptive area pattern of important comparative value. For leeches, while Schliep (1914) and Schmidt (1917, 1925a) provide the data which illustrate the way in which the presumptive area cells become segregated, their assignation to the fates listed above rests with the work of Whitman (1878, 1887), Nusbaum (1884, 1885, 1886), Biirger (1891, 1902), and Bychowsky (1921) on Glossiphonia and of Schmidt (1917) on Theromyzon, all of whom described aspects of later development. Presumptive mesoderm is segregated precociously in leeches into the cell 3D (Figs 12, 18), but the only significant difference between this and the polychaete-oligochaete condition is that presumptive midgut is eliminated from the D-quadrant. 3D divides equally bilaterally into M; and M, (Figs 19-21), teloblasts which, as in primitive oligo- chaetes, bud off paired ventrolateral mesodermal bands. The presumptive midgut is segregated, almost as in oligochaetes, into 3A, 3B and 3C (Figs 19, 20) of which the latter cuts off a small ectoderm cell 4c into the micromere cap before assuming its definitive fate. The position of the presumptive stomodaeum (Figs 18, 20) indicates that a major contribution is made to it by the anterior cells of the micromere cap, 2b and 3b, asin Tubifex. The presumptive embryonic ectoderm comprises a large dorsal posterior cell and the major part of the micromere cap in front of it. The large cell, as in Tubifex, is the main product of 2d and undergoes precocious division in the same sequence, equally bilaterally into four ectoteloblasts on each side (Figs 19-21) which together incorporate presumptive material of the prostomium, cerebral ganglion and the major part of the primordial epithelium of the trunk segments, and begin to bud off eight rows of small cells forwards, four on each side, as ectodermal bands. The micromeres multiply and spread back, between and around the ectoteloblasts and their products {Fig. 21), as in T'ubifex, forming presumptive temporary yolk sac, later to transform into segmental and pygidial epithelium. There is thus no doubt that cleavage and presumptive area formation in primitive oligochaetes and primitive leeches are fundamentally identical, supporting the classification of the two together as a single group, Clitellata. They can be interpreted as a particular modification of cleavage and presumptive area formation in polychaetes, associated with enhanced lecithotrophy and direct D. T. ANDERSON 19 embryonic development in a protective cocoon until numerous segments are formed and the adult organization is well established. It will be shown below that it is possible to argue in favour of the derivation of cleavage and presumptive area formation in Onychophora directly from a clitellate-like condition, assuming only further adaptation to increased lecithotrophy. Before pursuing this, however, the stability of presumptive area formation, in spite of secondary egg and cleavage specializations, will be displayed within the clitellates themselves by reference to the earthworms and to gnathobdellid and pharyngobdellid leeches. Cleavage and presumptive areas in lumbricid oligochaetes In earthworms, egg size is secondarily reduced, cieavage is modified and little trace of the spiral pattern remains. Attempts to draw comparisons with primitive oligochaetes and with polychaetes by means of the Wilsonian Figs 18-21.—18, presumptive areas of the blastula of Glossiphonia and Theromyzon. Diagram based on data from authors listed in the text. 19, Glossiphonia, postero-dorsal view of stage with paired M-cells. 20, Theromyzon, dorsal view of stage during ectoteloblast formation. 21, Glos stiphonia, postero-dorsal view after onset of ectoteloblast activity. (After Schliep, 1914 ; Schmidt, 1917, 1925a, 1944.) enumeration have therefore failed. The results obtained by Swetloff (1923a, 1928) on cleavage in Bimastus and Hisenia, using this notation, served at the time only to confirm the secondary specialization of earthworm cleavage, the bane of earlier workers (Wilson, 1889; Vedjovsky, 1888-92), without elucidating the mode of this specialization. The same results re-examined today, however, tell a different story, since they show that in earthworms the presumptive areas 20 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA of the blastula fall into a pattern identical with that of primitive oligochaetes, although the manner in which they are segregated is new and an additional paedogenetic structure has arisen. Taking Bimastus as an example, cleavage of the microlecithal egg is modified from the very beginning. The 1st division is unequal, yielding a smaller anterior and larger posterior cell (Fig. 22). At the 2nd division, the anterior cell divides equally into right and left daughters, while the posterior cell divides into a larger dorsal and a smaller ventral daughter (Fig. 23). The right anterior cell now divides again, completing a group of three anterior cells which undergo no subsequent division (Fig. 25). These three cells, which are characteristic of earthworms (Hatschek, 1878; Vedjovsky, 1887, 1888-92; Wilson, 1889 ; Hoffmann, 1899; Swetloff, 1923a, 1928) and finally become internal at the embryonic cell albumenotrophi 24 25 midgut Figs 22-25.—Bimastus. 22, 2-cell stage, dorsal view with posterior end on the right. 23, 4-cell stage, anterior view. 24, 7-cell stage, posterior view. 25, 10-cell stage, anterior view. (After Swetloff, 1923a.) anterior end of the embryo before being resorbed, are generally called provisional embryonic excretory cells. They become highly specialized, developing a complex series of intracellular canals, and show puisatory activity. There is no experimental proof of their excretory function, however, and it is equally possible that they act as absorption and assimilation of albumen until the embryonic cut cells assume this function. Meanwhile, continuing from the 4-cell stage, the posterior dorsal cell throws off a small cell downwards to the right, while the ventral cell cuts off a small cell upwards to the left (Fig. 24). The latter divides equally to form two small cells on the left, while the ventral cell cuts off a second small cell to the right (Fig. 25). The large dorsal cell now divides transversely into a smaller anterodorsal and a larger posterior cell. The latter then throws off a small posteroventral cell, somewhat displaced to the right. At the same time, the ventral cell has D. T. ANDERSON 21 divided equally twice, and its products, together with the posteroventral cell, make up a group of five ventral cells containing most of the yolk of the original egg (Fig. 27). These are the presumptive midgut cells. In front of them lie the three excretory cells. The large posterior cell divides equally bilaterally into two M-cells (Fig. 27), which subsequently bud off ventrolateral mesodermal bands. As in some polychaetes (Anderson, 1966a), the M-cells bud off one or two cells added to the presumptive midgut before assuming their definitive fate as mesodermal teloblasts. The anterodorsal sister of the M-cell mother cell divides equally bilaterally into two cells (Figs 26, 27) which, by further division, give rise to four ecto- teloblast cells on each side. The small cells on the right and left multiply and ectoteloblast cells yolk sac albumenotrophic midgut cells PL midgut yolk sac ectoderm mesoderm Figs 26—28.—Bimastus. 26, early blastula, anterior view. 27, early blastila, posterior view 28, presumptive areas of the blastula. (Based on data of Swetloff, 1923qa.) spread aS a micromere are in front of the anterodorsal cell (Fig. 26). In this are, the anterior cells in the mid-line are presumptive stomodaeum, the remainder temporary yolk sac. When cleavage in Bimastus is described in this way, it becomes obvious that the presumptive areas of the blastula (Fig. 28), although segregated by a new series of divisions consequent on secondary yolk loss, are identical in arrangement with those of primitive oligochaetes, the anterior cells having become paedo- genetically specialized without disturbing the configuration of the basic pattern. Presumptive midgut is relatively less extensive, commensurate with a reduction in yolk, but is still segregated into a group of cells ventrally. Presumptive mesoderm is contained within a pair of M-cells posteriorly. Presumptive ectoderm dorsally has a large central ectoteloblast mother cell surrounded 92 BWARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA anteriorly and laterally by an are of small cells. Presumptive stomodaeum is confined to the anterior members of this are, in the midline. The remainder develop as a temporary surface epithelium of the early embryo. Judging from the position of the excretory cells, lying between presumptive stomodaeum and midgut, it seems likely that they are derived from ancestral presumptive midgut cells, in which case an albumen-absorbing function becomes even more probable. Cleavage and presumptive areas in gnathobdellid and pharyngobdellid leeches Like earthworms, the gnathobdellid and pharyngobdellid leeches lay small, microlecithal eggs and show developmental modifications associated with prolonged feeding from an early stage on cocoon albumen (Schmidt, 1944). Cleavage among these forms has been studied only in Hrpobdella atomaria by Sukatschoff (1903) and Dimpker (1917) and in Hirudo medicinalis by Schoumkine (1953), but the results for the two species are sufficiently alike to suggest a basis of generalization for all gnathobdellids and pharyngobdellids. Not the least interesting aspect of cleavage and presumptive area formation in these animals is its similarity to that of earthworms, evolved in parallel from a fundamentally similar starting point. In spite of great reduction in egg size, the first three cleavage divisions retain the primitive hirudinean pattern. The Ist division is unequal, giving a larger CD cell and smaller AB cell, the 2nd division unequal in the former, equal in the latter, with the result that a relatively large posterior D cell is flanked by equal A, B and C cells, on the left, anteriorly and on the right respectively. At the 3rd division, a 1st quartette of micromeres is cut off dexiotropically to occupy a dorsal position, bilaterally arranged on either side of the mid-line, 1d and la to the left, 1b and le to the right (Fig. 29). Further divisions are now specialized (Fig. 30). The large posterior 1D cell first divides transversely and equally into posterodorsal 2d and postero- ventral 2D cells. The adjacent cell on the right, 1C, cuts off a small cell, 2c, upwards and inwards beneath the 1st quartette. 1A, 1B and 2C now do not divide again. Occupying the anterior and ventral surfaces of the embryo during cleavage, they eventually become overgrown during gastrulation and come to lie internally where they gradually degenerate. These cells clearly recall the three provisional excretory cells of earthworms. Although no mention is made in the work of Sukatschoff, Dimpker or Schoumkine of any pulsatory activity by the cells, or of the development of intracellular canals within them, it seems likely that their function is similar in leeches and oligochaetes, though whether excretory or absorptive 1s not yet clear. In leeches, as in earthworms, the relationship of the cells to the other presumptive areas suggests derivation from presumptive midgut, favouring a nutritive function. From the posteroventral 2D stem cell, a second 3d, and third 4d, cell are cut off into the interior, leaving the large cell 4D still at the surface. Meanwhile, the posterodorsal 2d cell cuts off two small cells 2d! and 2d*4 forwards at the surface behind the 1st quartette, leaving the large cell 2d?? still posterodorsal. The dorsal micromere cap now begins to multiply and spread (Figs 31, 32). Behind it, the large 2d cell divides equally bilaterally into two, each of which further divides into a typical row of four ectoteloblasts on either side of the dorsal mid-line (Figs 31, 32). Posteroventrally, the large 4D cell also divides equally bilaterally into two, each of which then divides into a lateral cell and a group of three medfan cells (Fig. 32). The latter push forward into the interior beneath the ectoteloblasts and make contact with the cells previously cut off into the interior from 10 and 2D. The lateral products of the 2D cell are the M-cells, which now begin to bud off mesodermal bands in the usual manner. The internal products of 2D, together with 2c, constitute the presumptive midgut. Taking into account the above series of divisions and what is known of gastrulation and subsequent development in Frpobdella and Hirudo, the D. T. ANDERSON 23 presumptive areas of these embryos (Fig. 33) can be seen to be identical with those of primitive leeches, if allowance is made for the paedogenetic specialization of the anteroventral and lateroventral stem cells and for a precocious movement of presumptive midgut cells into the interior during cleavage. The presumptive stomodaeum Figs 29-33.—EHrpobdella. 29, 8-cell stage, dorsal view with posterior end on the right. 30. 14-cell stage, left lateral view. 31, 16-cell stage, posterodorsal view. 32, 27-cell stage, posterior view. (After Sukatschoff, 1903; Dimpker, 1917.) 33, presumptive areas of Hrpobdella, based on data of Sukatschoff (1903) and Dimpker (1917). midgut, in association with secondary yolk reduction, is also much less voluminous than in primitive leeches, but is still segregated directly as a group of cells postero- ventrally. Presumptive mesoderm lies as a pair of M-cells, one on either side posterolaterally, bordering the posteriorly displaced presumptive midgut. Presumptive ectoderm, dorsally, has a large ectoteloblast mother cell behind a 24 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA group of micromeres. Presumptive stomodaeum is made up by cells at the anterior margin of this group. Furthermore, the parallel with earthworms as a similar response to a similar nutritional situation is unmistakable, and is especially emphasized by the paedogenetic specialization of the ‘‘ anterior ”’ presumptive midgut cells as provisional embryonic, possibly albumenotrophic cells. In spite of great modification of the cleavage sequence, differently in earthworms and in gnathobdellid and pharyngobdellid leeches, and of changed embryonic nutritional relationships, the basic presumptive areas remain constant in composition, relative juxtaposition and fate. Cleavage and presumptive areas in other clitellates In other clitellates morphologically more primitive than the earthworms and gnathobdellid and pharyngobdellid leeches, namely, the naidid oligochaetes and piscicolid leeches, the eggs are also microlecithal. Here, however, in spite of the systematic position accorded to these families in their respective subclasses, cleavage and the trophic relationships of the embryo in the cocoon are yet more specialized. As the work of Swetloff (1926) and Dawydoff (1942) on naidids and of Schmidt (1921, 1924, 1925b, 1930, 1939, 1941, 1944) on piscicolids shows, one of the products of cleavage is a provisional cellular envelope within which ectoteloblasts, M-cells and presumptive midgut become established and express their fate in a normal clitellate manner, and it seems highly probable that the provisional envelope has an absorptive function, rapidly supplemented by further, precociously formed temporary organs subserving feeding on the ambient nutrient albumen of the cocoon. Although it is not intended at the present time to discuss the development of these embryos in detail, there is no doubt that they, too, can be interpreted as modifications of primitive clitellate development in which the basic pattern of presumptive areas is retained in combination with adaptations to albumenotrophy, and they are mentioned here for two reasons. Firstly, they illustrate how even the most aberrant modifications, seemingly totally unintelligible when described by the Wilsonian system of enumeration, can be understood relative to their primitive antecedents when the formation and fate of presumptive areas is recognized. Secondly, like the earthworms and the gnathobdellid and pharyngobdellid leeches, they exemplify the fact that parallel developmental modifications can originate from parallel, related antecedents and need not necessarily be assigned to a common ancestry. Cleavage in the Onychophora As has been shown above, cleavage and presumptive area formation in primitive clitellates are adapted to direct lecithotrophic development within a protective cocoon, all known eases of the development of clitellates with small eggs being interpretable as instances of secondary yolk loss, associated with specialized albumenotrophy. Furthermore, lecithotrophic clitellate development manifests a particular modification of the pattern of total spiral cleavage and presumptive area formation seen in yolky polychaete embryos. While it is not possible on this evidence to single out any living polychaete family as a clitellate ancestor, the general notion of a monophyletic relationship between clitellates and polychaetes is supported by the evidence of development. Itis also clear that, as in polychaetes, the structure of the presumptive areas making up the wall of the blastula depends on the amount of reserve material incorporated into the egg and that cleavage is modified in ways commensurate with the establishment of these areas. Now, just as tlre primitive eggs of clitellate annelids are relatively large and yolky, so too are those of Onychophora. As has been demonstrated especially by Manton (1949), all instances of the development of small eggs in Onychophora reveal evidence of secondary yolk loss. Yolky onychophoran eggs, however, are larger than those of clitellates and individually enclosed in adherent egg membranes, and secondary yolk loss is associated, not with cocoon life, but with viviparity. Concomitantly, the early development of yolky eggs in Onychophora D. T. ANDERSON 25 is more highly modified than that of clitellates, the specializations associated with onychophoran viviparity are different from those associated with clitellate albumenotrophy, and the early embryology of the two groups seems quite dissimilar. chorion vitelline membrane cytopla ytopiasm 35 34 nucleus \ Ady any a a Ve, Wns ot = = = x vacuolated cytoplasm 36 oviducal wall oviducal epithelium egg Figs 34-37.—34, diagrammatic longitudinal section through yolky Onychophoran egg. 35, egg of Peripatopsis balfourt after dilatation. (After Manton, 1949.) 36, the same egg in transverse section. (After Manton, 1949.) 37, diagrammatic longitudinal section through upper oviduct of placental species, with zygote in lumen. Only a little is known of the yolky eggs of Onychophora, but it is clear that they combine dimensions of more than 1mm. (Ooperipatus, 1-9 mm., Dendy, 1902; Peripatoides novae zealandiae, 1:5mm., Sheldon, 1888; Peripatoides orientalis, 1-3mm., Anderson, unpublished; Hoperipatus weldoni, 1-3 mm., 26 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA Evans, 1902) with a primitive centrolecithal structure, dense with yolk, devoid of periplasm and containing the zygote nucleus in a more or less central cyto- plasmic halo (Fig. 34). Two membranes enclose the egg, a thin vitelline membrane and an external chorion which is thick in the oviparous Ooperipatus, thin in the ovoviviparous Peripatoides and Hoperipatus. The accounts of cleavage given by Sheldon (1888, 1889a) and oer (1902), although incomplete, show that in yolky onychophoran eggs it follows the typical arthropod mode, with nuclei and associated cytoplasmic haloes dividing and nucleus blastoderm 40 Figs 38-40.—Yolky onychophoran egg. 38, early cleavage, diagrammatic longitudinal section. 39, emergence of blastomeres at the surface, diagrammatic longitudinal section. 40, blastoderm stage, diagrammatic longitudinal section. increasing in number in the yolk, then rising to the surface (Figs 38, 39). At the same time, the yolk divides into a number of irregular spheres. Grouped. at first as a small disc of blastomeres, the superficial cells spread, with further divisions, to form a blastoderm enclosing the yolk spheres (Fig. 40). According to Sheldon, some of the nuclei and their cytoplasmic haloes in Peripatoides novae zealandiae remain within the yolk, but nothing is known of their fate, and Evans (1902) was unable to trace them in Hoperipatus weldoni. In general, it appears that cleavage in yolky onychophoran eggs is of a simple centrolecithal type, setting out a blastoderm around a yolk mass. T. ANDERSON 2h Development in the secondarily yolkless, viviparous Onychophora follows two modes, non-placental and placental. In the former, described with especial clarity for Peripatopsis by Manton (1949), the eggs released from the ovary are small and spherical (P. sedgewicki, 65-80u ; P. moseleyi, 150-172; P. balfouri, 380u; P. capensis, 260u; Sheldon, 1889); Bouvier, 1904; Manton, 1949), but on entry into the oviduct, they swell rapidly and become ellipsoidal (P. balfouri, Fig. 35) or cylindrical with hemispherical ends. Their dimensions are then, P. sedgewicki, 260 x 80u.; P. moseleyi, 520 x 160; P. balfourt, 480 x 220u; and P. capensis, 600 x145u (Manton, 1949) and the swelling can be interpreted as a delayed partial recapitulation of the enlargement which occurs in a yolky onychophoran egg during vitellogenesis. After swelling has taken place, the zygote nucleus lies in an irregular, granular cytoplasmic mass at one side of the egg (Fig. 36). The egg now becomes enclosed in two thin membranes which Manton (1949) suggests are vitelline membrane and chorion. Early cleavage is similar in all four species (Figs 41-47). The cytoplasm of the egg breaks up into a number of spheres of different sizes, recalling the yolk spheres of Peripatoides and Hoperipatus, floating freely inside the egg membranes in a watery fluid. The zygote nucleus lies in one of the spheres, while the remainder are anucleate pseudoblastomeres. The onset of cleavage mitoses is thus somewhat delayed. When it occurs, the 1st blastomere divides to form a disc of blastomeres apposed to the inner surface of the egg membranes on one side of the egg, recalling the blastomere dise of the yolky-egged species. As they form, the blastomeres of the disc gradually absorb the material of the disintegrating pseudoblastomeres. By the time 64 cells are present, the blastomere disc extends as a saddle around two-thirds of the egg circumference, leaving one side and both ends blastomere-free. In P. sedgewicki and P. moseleyi, the saddle of blastomeres now gives rise by further cell division and marginal closure to a blastoderm of about 160 cells around a central fluid-filled space (Fig. 48). Cell division continues and at the same time the vitelline membrane is absorbed and the blastoderm swells by fluid uptake, stretching the outer membrane, until its diameter becomes, in P. sedgewicki, about 1000 500u, in P. moseleyi, about 900 x500u. The only difference between these embryos and the more primitive yolky onychophoran embryos at the same stage is replacement of yolk by fluid in the blastocoel. In P. capensis, cleavage and blastoderm formation overlap a precocious onset of movement of cells into the interior (Figs 49, 50). At the edges of the saddle of blastomeres, cells enlarge and move into the concavity of the saddle This process continues as the saddle cells divide further and spread as a blastoderm, so that the inner surface of the blastoderm becomes lined with large, vacuolated cells. The edges of the outer layer finally form a slit orientated along the anteroposterior axis of the embryo, and with closure of the slit the blastoderm becomes complete. No dilatation of the blastoderm occurs in P. capensis, although the surrounding egg membranes swell to the expected size as a result of fluid ingress. In P. balfouwrt (Figs 51-54), the marginal cells of the saddle of blastomeres also enlarge and move into the interior as the saddle spreads to form a blastoderm, so that the blastocoel is temporarily filled with a mass of vacuolated cells ; but these now disintegrate, while at the same time the blastoderm dilates, attaining a final condition like that of P. sedgewicki and P. moseleyi. The significance of the cleavage peculiarities of P. capensis and P. balfouri as compared with P. sedgewicki and P. moseleyi will be taken up below. The placental viviparous Onychophora have eggs which are even smaller than those of the non-placental species and, furthermore, lack the recapitulatory swelling of the egg after release from the ovary (Fig. 37). Dimensions lie between 25u and 40u (Bouvier, 1904) and enclosing membranes are absent (Kennel, 1884, 1885; Sclater, 1888). Cleavage in these eggs is little understood. Kennel (1884, 1886) and Sclater (1888) showed, and further unpublished observations 28 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA by Manton and myself have confirmed, that it is total and equal, resulting in the formation of a morula lying in the lumen of the oviduct (Fig. 55). With further cell divisions, the morula becomes hollow, with a wall one cell thick, before attaching to the oviducal wall (Fig. 56). Rather than dilating to form a blastoderm, the hollow vesicle enters immediately into a further specialized phase, in which the cells in the region of attachment multiply to form a hollow stalk adpressed terminally as a flat placental plate against the oviducal wall, pSseudoblastomere ak as) me ed. Figs 41-43.—Peripatopsis moseleyi. 41, 2-cell stage. 42, 21-cell stage. 43, saddle of blastomeres (After Manton, 1949.) Figs 44-47.—Peripatopsis balfowri. 44, 1-cell stage. 45, 8-cell stage. 46, 1-cell stage, transverse section. 47, 16-cell stage, transverse section. (After Manton, 1949.) D. T. ANDERSON 29 while the remaining cells multiply to increase the size and cell number of the hollow vesicle at the free end of the stalk (Fig. 57). The further development and comparative significance of the stalk and vesicle will be examined below. - Developmental fate of the cleavage blastomeres in Onychophora Just as in the Clitellata, the cells of the blastoderm or its equivalent immediate pre-gastrula stage in Onychophora can be assigned to presumptive areas of specific subsequent fate by taking account of the events of gastrulation and later development. The focal point in the elucidation of these presumptive areas is the composition of the blastoderm in yolky onychophoran embryos. Unfortunately, one of the weaknesses of Sheldon’s work on Peripatoides novae saddle of SSS SELL ep eae \94 OE Cal Figs 48—50.—48, Peripatopsis moseleyi, blastoderm stage before dilatation. 49, P. capensis, section through late cleavage stage. 50, P. capensis, section through almost completed blastoderm. (After Manton, 1949.) zealandiae, and a matter calling urgently for reinvestigation, is a lack of accurate description of gastrulation and early segment formation. At the same time, Sheldon’s results, together with those of Evans (1902) and some preliminary unpublished observations of my own on embryos of Peripatoides orientalis from New South Wales, permit the tentative construction of a presumptive area map for ovoviviparous species (Fig. 58). Anteroposterior and dorsoventral axes are fixed, the former corresponding to the long axis of the egg. Midventrally lies a long narrow band of presumptive midgut cells, soon to migrate into the interior, leaving an elongate slit at the surface. At the margin of the presumptive midgut lie, anteriorly, presumptive stomodaeum, laterally, paired ventrolateral bands of temporary yolk sac ectoderm and, posteriorly, presumptive proctodaeum. Midventrally behind the latter lie small areas of presumptive posterior midgut and presumptive mesoderm. Associated with the latter are presumptive germ 30 EARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA cells. Lateral to the ventral areas listed above, and converging posteriorly on the presumptive mesoderm, are two broad bands of presumptive ectoderm. Dorsally lies a further broad area of temporary yolk sac ectoderm. The data of Manton (1949) permit much more firmly substantiated presumptive area maps to be drawn for the viviparous non-placental Peripatopsis. In no case in this genus does the anteroposterior axis of the embryo correspond vestigial anterior midgut Figs 51—-54.—Peripatopsis balfourt. 51, saddle of blastomeres stage. 52, same embryo from opposite side, showing vacuolated cells. 53, transverse section through early saddle of blastomeres stage. 54, transverse section through blastoderm stage. (After Manton, 1949.) to the long axis of the egg. It may fall in any direction, suggesting that the axial relations of the embryos are established late, after blastoderm formation, by processes which are at present obscure. This, however, does not confound the reality of the presumptive areas which can be discerned. In P. sedgewicki they centre on a terminal, and in P. moseleyi a lateral cell group which lie postero- ventrally relative to future development. The cells of the group (Fig. 59) D. T. ANDERSON 31 comprise presumptive mesoderm, presumptive posterior midgut anterior and internal to the mesoderm, presumptive ectoderm marginally, and a number of presumptive germ cells. The latter tend to lie between the presumptive midgut and presumptive mesoderm in P. moseleyi and at the posterior end of the pre- sumptive mesoderm in P. sedgewicki, a minor discrepancy of localization with no special significance. Immediately anterior to the presumptive posterior midgut lie the presumptive proctadaeum and stomodaeum, conjoined. Lateral to them are ventrolateral bands of presumptive temporary yolk sac ectoderm, followed by oviducal epithelium cleavage morula 56 blastodermic OQ placental plate vesicle SOQWIH KOS? S ze © LS O96 0 ae as Racca Oy EG ee cop per Wa SS a U Z een K.\ FO ORS AZ 6 Ay Kors 57 placental stalk Figs 55-57.—Viviparous placental onychophoran. 55, early cleavage morula. 26, later morula attached to oviducal wall. 57, blastodermic vesicle and placental stalk. broad bands of presumptive embryonic ectoderm, with the remainder of the blastoderm, laterally and dorsally, constituting further temporary yolk sac ectoderm. The blastodermal presumptive areas of P. balfouri are identical with those described above, with the presumptive germ cells arising from the presumptive mesoderm as in P. sedgewicki. It is necessary to take into account, however, the temporary vacuolated cells budded off in P. balfourt from the edge of the saddle of blastomeres into the interior during blastoderm formation. Since the point towards which these proliferating edges converge and close subsequently forms the presumptive stomodaeum and proctodaeum, a place can be assigned 32. BARLY EMBRYOLOGY OF OLIGOCHAETA, HIRUDINEA AND ONYCHOPHORA along the ventral midline of the latter as the site of origin of the temporary vacuolated cells, already budded off. By comparison with the presumptive area map for yolky species, the temporary vacuolated cells would appear to be modified midgut cells, as suggested by Manton (1949). ectoderm 7 yolk sac ectoderm posterior midgut ectodermal mesoderm 58 growth zone 59 posterior midgu mesoderm stomo-proctodaeum mesoderm 60 ectoderm stomodaeu Calogyne (2-fid) Fig. 10. Diagram to show the suggested relationships of the types of seed found in Goodenia and its satellite genera. Hatched areas=mucilage cells; solid black areas=thickened epidermal cells ; stippled areas=endosperm and embryo. R. CG. CAROLIN 79 Yet another line is shown by type 6, in which one of the marginal thickened cells becomes much elongated and supports the large ‘“‘ mucilage ”’ cells overlying the outer parts of the body of the seed. These peculiarities are relatively undeveloped in G. pterygosperma and G. trichophylla but extremely well developed in G. micrantha. Type 7 is a very reduced form and it is difficult to fit it into any of these lines; it could be the end product of reduction of any of them. Without evidence from other characters, which is being accumulated, it is impossible to place this species in any series. These lines all seem to converge on that of type 2 (a) and, to some extent, intergrade with it. This is further evidence for considering it to be the basic, primitive type. Despite these intergrades the types are fairly distinct and close-knit and, for this reason, represent real biological units. Evidence is being accumulated that other distinctive characters are correlated with them. This being so it would appear that Krause’s sections and series (1912) require a drastic overhaul. Of his sections, Monochila is particularly unsatisfactory, since it contains some species having type 2 and one species with type 1 seeds. G. barbata, G. strophiolata (and G. chisholmit), from section Hugoodenia should be linked with G. phylicoides. Ser. Suffruticosae seems to be a rather hetero- geneous group and needs breaking up. Ser. Caeruleae divides quite neatly into two groups with type 2 and type 6 seeds respectively. Ser. Racemosae is fairly uniformly type 3. Ser. Foliosae, on the other hand, splits into three fairly well defined groups: those centred around G. grandiflora with type 2 (b), those around G. hispida with type 3, and those around G. heterochila with types 4 and 5. There seems little doubt that this is a heterogeneous taxon when such widely differing seed types are found within it. It is interesting to find that G. grandiflora frequently has bracteoles despite Krause’s placing them in the Ebracteolatae. Indeed, the presence or absence of bracteoles seems to be of little value in itself. To turn to the ‘satellite’ genera. Calogyne again has shown its heterogeneous nature (see Carolin, 1959). C. berardiana belongs to the evolutionary line represented by G. glauca and G. heterochila. All the other species belong to the line represented by G. hispida and G. koningsbergert. Calogyne, as constituted, is an unsatisfactory genus and the 2-fid-styled species should be separated from those having 3-fid styles. Selliera belongs to the evolutionary line of type 3 similar to G. koningsbergeri which, indeed, has been referred to Selliera in the past. Neogoodenia has a rather more basic seed-type, and it is consequently rather more difficult to assign it to any particular line of development. Although there is only a single seed in an indehiscent fruit, there is no doubt of its close affinity with Goodenia on account of its seed type. All the “ satellite’ genera so far discussed are so close to Goodenia that a case could be made for their inclusion in that genus. The seeds of Velleia are very similar to types 3(b) and 2(b) of Goodenia. Thus, although they possess the ‘‘ mucilage”’ cells, and thus a differentiated seed coat, the type shown is a fairly basic one. It seems probable, then, that Velleia has separated from the bulk of Goodenia subsequent to the differentiation of the mucilage cells in phylogenetic history, and thus after both had diverged from the G. barbata group: a further argument for accepting the latter as a separate genus. Velleia, however, must remain a generic entity by virtue of its other characters. Both Catosperma and Pentaptilon show a seed structure undoubtedly derived from a Goodenia type, but the reduction which has apparently accompanied the closure of the fruit has made it impossible to decide to which line they belong on this evidence. 30 SEEDS AND FRUIT OF THE GOODENIACEAE Verreauaia differs from all the other genera examined in this group so far by its lack of mucilage cells. All the epidermal cells of the compressed seed, even those on the margins, are uniformly thickened. They are distinctly thickened and the vasculature is quite definitely of the Goodenia type—there is no doubt about the position of the genus in this group. Thus Verreauxia is either a case of extreme reduction, the possibility of which is shown by Goodenia koningsbergert, or has retained the primitive condition also shown by the G. barbata group. Evidence from other directions is needed before this problem can be resolved. The discovery of the 4-locular nature of the ovary of Scaevola porocarya is of some significance in that it forces a new interpretation of the ovary structure, at least of Scaevola. Figures 9, A-G, show that the loculi must be considered as carpels; they have a vasculature in fair agreement with this hypothesis, although those of the opposing pairs of carpels have reacted in different ways to the adnation of the outer whorls to the ovary. The lateral pair of carpels have a dorsal strand each, produced from the union of two strands, themselves derived separately from the central stele. This is a very unusual condition ; although the double nature of the dorsal bundle has been established in some primitive groups it is unexpected here. The posterior-anterior pair have a dorsal vasculature exactly similar to that found in other Scaevola spp. (Carelin, 1959). The adjacent lateral bundles of the carpels are united and the laterally opposite pairs eventually unite. Neither these nor the dorsal bundles of the lateral carpels extend into the style, rather unusual, as in other species the former do so and in S. ramosissima all four lateral bundles separate and are persistent through the style (Carolin, 1959). It appears that the evolution of the vascular system of the flower is just as uneven as the larger differences found in the Dampiera group (see above), advanced and primitive conditions occurring in the same species, indeed, at different levels within the same flower. It seems, then, that Scaevola has had a rather different history from that of the rest of the Goodenia group. The behaviour of the “ pseudo-carpellary dorsal ”’ bundles at the top of the ovary of Goodenia, etc., implies that the loculus is derived from two carpels (Carolin, 1959). The bundles similarly positioned in Scaevola remain single always; this fact, together with the discovery of the structure of S. porocarya, implies that the loculus in the case of Scaevola represents a single carpel. Furthermore, if this reasoning is accepted, Goodenia has ‘‘ true ” carpels placed diagonally in the flower whilst Scaevola has them placed anterior- posteriorly and (sometimes) laterally. This is not necessarily a particularly far-reaching difference since these two conditions may occur elsewhere in the plant kingdom within the same species (e.g., Campanula). It does, however, imply that the evolutionary histories of Scaevola and Goodenia are separate, and similarities in the ovary structure are the result of convergence rather than common origin. The seeds of Scaevola are usually more or less compressed, although the structure of the testa is rather similar to that of Dampiera. However, there is little doubt that it is not closely allied to Dampiera since the seed has a single strand, the floral structure is different (Carolin, 1959) and the chromosome number is different (Peacock, 1961). Within the genus the fruit types (see Fig. 11), once again, show only incomplete correlation with the Sections proposed by Krause. After the type exhibited by S. porocarya, it seems reasonable to suppose that type 2 is the most primitive since, in this case, the tissues which are most probably gynoecial in origin behave as a unit, i.e., form the ‘ endocarp’’. The false-loculi really represent a case of ‘‘ regressive convergence’ towards the condition in type 1 subsequent to the loss of the lateral carpel (loculi). These taxa belong mostly to the Sarcocarpaea and all of them have a highly developed “‘ mesocarp’’, be it fleshy or corky. The trend in the Sections Crossotoma, Xerocarpaea and Pogonanthera is towards the exclusion of part of the tissue of gynoecial origin R. C. CAROLIN 81 from ‘‘ endocarp’”’ formation, this part being around the false-loculi caused by the disruption of the “ lateral’? bundles. In sect. Crossotoma no false- loculi are formed and these tissues are included in the fleshy ‘‘ mesocarp ” ; in sects Xerocarpaea and Pogonanthera the false-loculi are frequently present and the ‘‘ mesocarp ”’ is thin. S. crassifolia and S. nitida resemble S. porocarya 7 Fig. 11. Diagram to show the suggested relationships of the pseudo-fruit types of Scaevola. Open areas=dry, soft tissue; solid black areas=dry, hard tissue; stippled areas=fleshy (or corky) tissue. so closely in many respects there seems little doubt that they are related despite the first two species having type 2 fruits. It appears, then, that the Sarcocarpaea have originated from the 4-locular type separately from the Xerocarpaea group and possibly earlier. Sect. Pogonanthera shows very few differences from Xerocarpaea in fruit structure and sect. Crossotoma appears to be a fleshy- ‘““mesocarp ’’ derivative of the Xerocarpaea. F 82 SEEDS AND FRUIT OF THE GOODENIACEAE The three species, S. stenophylla, S. helmsivi, S. fasciculata, show very real differences in the seed form from the rest of genus ; they also had a significantly different floral construction (Carolin, 1959). In fact, the seeds have very definitely thickened epidermal cells, quite unlike the lignin banding seen in the rest of Scaevola spp. Indeed, they resemble those of Verreauxia or Goodenia without the marginal mucilage cells. It was previously suggested that these species were closer to Verreauxia than Scaevola (Carolin, 1959) but other characters would seem to link them with a group of Goodenia spp., centred around G. scapigera. This would, in fact, imply not only an evolutionary trend towards reduction in ovule number and closure of the fruit, but also a loss of the mucilage cells and a reversal, in the evolutionary sense, to something more closely resembling the G. barbata type. The seed of S. helmsti, however, is compressed and not + circular in transection. The rest of Scaevola ser. Uniloculatae shows a straightforward unilocular drupaceous fruit with a thin “‘ mesocarp ”’ and no false-loculi. Although there is no indication from the fruit or seed that there have been several origins of the unilocular condition from the (apparently) bilocular condition (cf. Carolin, 1959), other characters ‘indicate that this may be so. Acknowledgements Grants from the University of Sydney and the Australian Research Grants Committee enabled this work to be completed. References BENTHAM, G., 1869.—Flora Australiensis, vol 4. London. Bersier, J. D., and BocqueEt, G., 1960.—Les méthodes d’éclairissements en vascularisation et en morphogenic végétales comparées. Arch. Sci. Phys. et Nat. (Gen.), 13: 555. Caro.in, R. C., 1959.—Floral Structure and Anatomy in the Family Goodeniaceae. Proc. Linn. Soc. N.S.W., 84: 242. , 1960.—The structures involved in the presentation of pollen to visiting insects in the Order Campanales. Proc. Linn. Soc. N.S.W., 85: 197. , 1960b.—Floral structure and anatomy in the Family Stylidiaceae. Proc. Linn. Soc. N.S.W., 85: 189. GARDNER, C., and GEORGE, A., 1963.—EHight new plants from Western Australia. Jowr. roy. Soc. W.A., 46: 129. Guick, D., 1949.—** Techniques of histo- and cytochemistry ”’. New York. Krause, K., 1912.—‘‘ Das Pflanzenreich ’’, 54—Goodeniaceae und Brunoniaceae. Berlin. Linpiey, J., 1836.—‘‘ Natural System of Botany ”’. London. Martin, A. C., 1946.—The comparative internal morphology of seeds. Am. Midl. Nat., 36: 513. Pracocr, W. J., 1963.—Chromosome numbers and cytoevolution in the Goodeniaceae. Proc. Linn. Soc. N.S.W., 88: 8. SKOTTSBERG, C., 1923.—Artemisia, Scaevola, Santalum and Vaccinum of Hawaii. Bull. Bish. Mus., 43: 1. SreEpMAN, H. F., 1960.—“ Section cutting in microscopy’. Oxford. Voucher Specimens Leschenaultia biloba Carolin 3428 (Syd.). JL. divaricata Carolin 43 (Syd.). L. filiformis Perry 1710 (Canb.). LZ. linarioides Carolin 3324 (Syd.). LZ. longiloba no voucher. Anthotium rubriflorum no voucher. Dampiera coronata Carolin 3243 (Syd.). D. purpurea Carolin 4439 (Syd.). D. stricta Peacock 6011.26.6 (Syd.). D. trigona Carolin 3055 (Syd.). Diaspasis filiformis no voucher. Goodenia affinis Carolin 3340 (Syd.). G. amplexans Black, Burnside Road 2.1904 (N.S.W.). G. armstrongiana Chippendale 18.5.59 (N.S.W. ex N.T. 6199). G. armitiana Perry 1904 (Canb.). G. auriculata Perry 1772 (Canb.). G. azwrea Chippendale 12.4.59 (N.S.W. ex N.T. 5659). G. barbata Peacock 6012.5.1 (Syd.). G. bellidifolia Carolin 565 (Syd.). G. caerulea Sargent 4069 (N.S.W.). G. boormanii Peacock 6111.26.1 (Syd.). G. calcarata Beadle 8.1941 (Syd.). G. chisholmit Peacock 6012.4.1 (Syd.). G. concinna Benn 4.10.1963 (Syd.). G. corynocarpa no voucher. G. decurrens Peacock 6011.26.3 (Syd.). G. dimorpha Carolin Oct. 1963 (Syd.). G. disperma Mueller, Burnett River (K). G. eatoniana Koch, 2032 (N.S.W.). G. glabra Peacock R. C. CAROLIN 83 6111.31.1 (Syd.). G. glawea Wharton 9.2.1952 (Syd.). G. grandiflora Cambage 3978 (Syd.). G. havilandii var. pawperata Specht and Carrodus 34 (Syd.). G. hederacea Carolin 2044 (Syd.). G. heterochila Perry 2405 (Canb.). G. heterophylla Carolin 2032 (Syd.). G. hispida Eddy, 23.3.58 (N.S.W. ex N.T. 5220). G. incana Cleland, Nargin, 11.08 (N.S.W.). G. koningsbergeri no voucher. G. laevis Carolin 3583 (Syd.). G. macmillanii Lucas, Macalister River, 12.10 (N.S.W.). G. micrantha Peacock 60856.5 (Syd.). G. microptera Chippendale 2.5.58 (N.S.W. ex N.T. 4232). G. mitchellii Carolin 4055 (Syd.). G. ovata Carolin 1936 (Syd.). G. paniculata Peacock 6012.2.2 (Syd.). G. phylicoides Carolin 3575 (Syd.). G. pinnatifida Peacock 6111.9.1 (Syd.). G. ~pterygosperma Bern 8.10.1963 (Syd.). G. pumilio McKee 9471 (N.S.W.). G. pusilliflora Whaite 2491 (N.S.W.). G. quadilocularis no voucher. G. ramellii Perry 1918 (Canb.). G. rotundifolia Peacock 6012.6.3 (Syd.). G. scaevolina Perry and Lizarides 2296 (Canb.). G. scapigera Carolin 3556 (Syd.). G. sepalosa Perry 1708 (Canb.). G. stelligera Peacock 6012.12.1 Syd.).) G. strophiolata Doodlakine, Fitzgerald Nov. 1903 (N.S.W.). G. tenwiloba Peacock 60865.1 (Syd.). G. trichophylla Pritzel 827 (N.S.W.). G. varia Maiden Kangarcalslaw 1902 (N.S.W.). G. viscida no voucher. G. vilmoriniae Perry and Lazarides 2225 (N.S.W.) Selliera radicans Peacock Jan. 1958 (Syd.). Calogyne pilosa Specht 218 (Canb.). C. berardiana Carolin 4224 (Syd.). Neogoodenia minutiflora George 910 (Perth). Symphiobasis macroplectra Speck 1368 (N.S.W.). Pentaptilon careyi Carolin 3325 (Syd.). Verreauxia reinwardtii Peacock 60845.1 (Syd.). V. paniculata no voucher. Catosperma goodenicearum Chippendale 14.7.1956 (N.S.W. ex N.T. 2331). Scaevola albida Carolin 2082 (Syd.). S. calendulacea Carolin 0785 (Syd.). S. chamissoniana Smith-White, Aug. 1961 (Syd.). S. crassifolia Carolin 3373 (Syd.). S. dielsii Briggs (N.S.W. 52429). SS. fasciculata Briggs (N.S.W. 52427) S. glandulifera Peacock 60876.1 (Syd.). S. globulifera Briggs 3.10.1960 (Syd.). S. helmsii Carolin 3134 (Syd.). S. hookert Carolin Jan. 1957 (Syd.). S. huméifusa Carolin 3538 (Syd.). S. indigofera McKee 4714 (Syd.). S. micrantha McKee 4279 (Syd.). S. mollis Smith- White, Aug. 1961 (Syd.). S. nitida Carolin 3190 (Syd.). S. ovalifolia Carolin 4270 (Syd.). S. oppositifolia no voucher. S. porocarya Carolin 3358 (Syd.). S. ramosissima Carolin 3653 (Syd.). S. spimescens Carolin 3074 (Syd.). S. taccada 8.U. Biol. Soc. Jan. 1948 (Syd.). S. tomentosa Carolin 3323 (Syd.). FE FUNGI ATTACKING SEEDS IN DRY SEED-BEDS | D. M. GRIFFIN School of Agriculture, University of Sydney, N.S.W., Australia [Read 27th April, 1966] Synopsis Soil samples were brought to approximate equilibrium with atmospheres of 100, 95, 90, 85 and 80% R.H. The pattern of fungal colonization of wheat seeds buried in the soil samples was then compared with that occurring in seeds stored at similar humidities, and was found to be similar. The breakdown of buried seeds was due to such seed-borne fungi as members of the Aspergillus glaucus group and the Penicillium chrysogenum and the P. citrinum series and to various soil-borne aspergilli and penicillia, of which P. expansum was the most important. Only in the 80% R.H. series did most of the buried seeds survive for more than three weeks and the period was less than one week for those in the 100% and 95% R.H. series. Shaking the seeds with “ New Improved Ceresan ” fungicidal dust prior to planting did not enhance longevity. INTRODUCTION The subject of deterioration of stored grain due to fungal activity has been reviewed by Christensen (1957) who concluded that fungal invasion was the main cause of damage to the embryos of cereals. Species of Aspergillus and Penicilium are able to cause seed decay on an economically-important scale in atmospheres with a relative humidity as low as 70% (Snow, Crichton and Wright, 1944; Snow, 1945). Although fungi are active in soil at similar low relative humidities (Dommergues, 1962; Griffin, 1963b; Kouyeas, 1964), there has been no study of germination reduction in seeds sown in soil with a moisture content below that of the permanent wilting point. In Australia and similar regions, however, this is a matter of some importance, for germination of seed is often retarded by low soil moisture at, or immediately after, sowing. MATERIALS AND METHODS To control relative humidity in the experiment using soil, thin layers of air-dry sieved soil were placed in small containers in each of six vacuum desiccators, the basal portions of which contained one of the six sodium chloride solutions listed in Table 1. The desiccators were then evacuated for one week to allow the soils to attain the required relative humidity. The water contents of soil samples (Table 1) indicate that the 100% R.H. sample was at approxi- mately permanent wilting point and that those from other humidities were considerably drier. Sixty seeds were placed in perforated plastic containers hanging above c. 170 ml. solutions of sodium chloride in closed jars of 500 ml. capacity. In the experiments using soil, about 100 g. of soil, after equilibration, were also placed in the containers. After 10 weeks, a test using Clark’s and Severinghaus’ electrodes showed an oxygen concentration in the jars not below 19% and a carbon dioxide concentration not above 2%. Any change in the atmosphere of the jars was therefore negligible. In a small pilot experiment, variation between replicates in both the pattern of colonization and the rate of seed degeneration was also found to be negligible and the main experiments were therefore not replicated. PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WAtEs, Vol. 91, Part 1 D. M. GRIFFIN 85 TaBLe I Moisture data for soil samples Satu- Field Perma- Relative humidity rated capa- nent soil city wilting 100 95 90 85 80 point Controlling salt solution 0 8-6 16-5 23-6 30-0 — —- — (g. NaC1/100 ml. H,O) Suction (pF)* .. Sy ASF 4-86 5-17 5:36 5-50 0 2:0 4-2 Moisture content (g. 21 10 9 8 ates 63 39 18 H,0/100 g. dry soil) Moisture content Je) 80 16 14 13 11-6 100 62 29 (% saturated) * Relationship between soil water suction and R.H. has been given by myself (Griffin, 1963a, 6) with an error in the formula. M=molecular weight of water, not density. + Not in true equilibrium. The soil was a black earth (Stephens, 1962) of very heavy texture (pH 8-6) from Narrabri in the north-west wheat areas of New South Wales. The wheat seeds were of the variety Mengavi and were harvested approximately one year before the start of the experiments. At each sampling, five seeds were taken from each relative humidity and were sterilized by successive immersions in (1) 95% alcohol (momentarily), (2) 1% silver nitrate (0-5 min.) and (3) saturated sodium chloride (3 min.). Each seed was then cut aseptically into four portions, three seeds being placed on Petri dishes containing Czapek-Dox agar (39% sucrose) and two on dishes containing Czapek-Dox Sucrose agar (20° sucrose). Seeds from the same containers were also tested for ability to germinate when placed on wet filter- paper in a Petri dish. All experiments were conducted at 25°C (+0-5°) in a dark room and no water was deposited on the surfaces of the containers. TABLE 2 Number of seeds germinating, out of ten, after various periods in controlled humidity jars Relative Humidity Week 100 95 90 85 80 1 10 10 10 10 10 2 3 6 8 9 10 3 2 2 5 7 10 4 1 3 2 a 8 5 2 0 4 3 10 7 0 2 1 2 9 9 0 0 3 2 6 13 ate 0 1 0 1 17 arb Eee ast ae 2 EXPERIMENTAL Internal microflora of seeds The experiment was made to study the pattern of colonization of the interior of the seed by the seed-borne flora. Wheat seeds alone were therefore placed in the experimental containers. 86 FUNGI ATTACKING SEEDS IN DRY SEED-BEDS Data on seed germination are given in Table 2. The fungi isolated from surface-sterilized seed were, with very few exceptions, species of Aspergillus and Penicillium. Owing to the presence of many intermediary forms, these fungi have not been identified at the species level but have been referred in Table 3 to the appropriate group or series (Thom and Raper, 1945; Raper and Thom, 1949). Within the important A. glaweus complex, A. amstelodami (Mang.) Thom and Raper, A. chevalieri (Mang.) Thom and Raper, A. repens (Cda.) De Bary and A. ruber (Spiek. and Brem.) Thom and Church were isolated. Other fungi occasionally isolated but not included in Table 3 were Alternaria sp., Aspergillus ochraceus Wilhelm, A. ustus (Bain.) Thom and Chureh, Mucor sp., Penicillium lilacinum Thom and P. purpurogenum Stoll. TABLE 3 Aspergilli and Penicillia isolaied from surface-sterilized seeds Relative Humidity Week 100 95 90 85 80 1 P. chrysogenum PP. chrysogenum P. chrysogenum 2 A. candidus A. glaucus P. chrysogenum P. chrysogenum FP. chrysogenum 3 A. glaucus P. chrysogenum A. glaucus P. citrinum 4 P. chrysogenum A. candidus A. glaucus A. glaucus A. glaucus P. chrysogenum P. chrysogenum P. citrinum 5 A. candidus A. versicolor A. candidus A. glaucus P. citrinum A. glaucus A. versicolor P. citrinum P. citrinum 7 A. versicolor A. glaucus A. glaucus A. flavus-oryzae A. glaucus A. versicolor A. glaucus P. citrinum P. chrysogenum 9 A. flavus-oryzae A. candidus A. fiavus-oryzae A. glaucus A. glaucus P. chrysogenum A. flavus-oryzae A. glaucus P. citrinum P. citrinum P. citrinum A. versicolor P. citrinum P. citrinum 13 — A. glaucus A. glaucus A. glaucus A. glaucus A. versicolor A. versicolor P. chrysogenum FP. citrinum P. chrysogenum P. citrinum P. citrinum 7 oo — — — A. glaucus P. chrysogenum Internal microflora of seeds in contact with soil Wheat seeds were buried in small quantities of soil in the experimental containers and the pattern of colonization of the interior of the seeds was again studied. ~ Data on germination are given in Table 4 and on the commonest fungi isolated (identified to group or series only) in Table 5. Other fungi sometimes isolated were Aspergillus avenaceus G. Smith, A. flavepes (Bain. and Sart.) Thom and Church, A. ochraceus Wilh., Cephalosporium sp., Fusarium spp., Penicillium brevi-compactum Dierckx, P. canescens Sopp, P. frequentans Westling, P. funiculosum Thom, P. oxalicwm Currie and Thom, P. purpurogenum Stoll, and Rhizopus stolonifer Fr. D. M. GRIFFIN 87 An attempt to improve longevity by coating seeds with a fungicidal dust before burying was unsuccessful. The seeds were shaken-up with an excess of ‘“New Improved Ceresan”’ dust (ethyl mercuric phosphate—1 -3°% Hg) and then buried, but the decrease in viability was very similar to that in the previous experiments and the fungicide had clearly been ineffective. The only fungus to be isolated from any of the seeds was Penicillium expansum Link and this species was shown in a subsidiary experiment to be unusually tolerant of the fungicide when it was added to agar media. On an agar medium containing ce. 1 p.p.m. of the fungicide, only P. expansum and P. citrinum, of the fungi tested, suffered no check to growth whereas the growth rates of P. chrysogenum, P. urticae, Aspergillus versicolor and A. ruber were, at most, half of the normal. P. expansum was the only fungus little affected by a concentration of ¢. 100 p.p.m. TABLE 4 Number of seeds germinating, out of ten, after various periods in soil in controlled humidity jars Relative Humidity Week 100 95 90 85 80 1 1 4 10 10 10 2 2 1 4 10 10 3 0 0 2 7 8 4 0 0 2 10 5 2 0 0 0 10 7 ae 1 0 0 7 9 == 0 2 3 4 13 = 0 0 3 0 DISCUSSION The pattern of fungal colonization of the unburied seeds is similar to that described by many previous workers studying the deterioration of stored seed, with members of the Aspergillus glaucus group and the Penicillium citrinum and P. chrysogenum series being especially prominent. The rate of colonization increased with increasing humidity and there was little fungal invasion at 80% R.H. until five weeks had elapsed. With the seed buried in soil, a greater variety of fungi was recorded, but the rate of deterioration was approximately the same as in stored seed. From a comparison of the fungi listed in Tables 3 and 5 it can be seen that seed destruction in soil was due to both seed-borne and soil-borne fungi, and of the latter, Penicillium expansum was particularly important. In both experiments, early records of a fungus at the lower humidities presumably indicate its presence as spores or a few hyphae in the more super- ficial layers of the seed but still in a position shielded from the effects of surface sterilization (Hyde, 1950, 1951), so that the presence of the fungus could not be detected visually until after incubation on agar. Dead seeds, however, usually bore an external mass of conidiophores and were also heavily colonized internally. It seems reasonable to suppose that this fungal invasion was the cause of death, for cereal seeds seem to offer no resistance to fungal attack when held at moisture levels sufficient for fungal growth but insufficient for seed germination. Such a contention is also supported by the fact that the rise in respiration rates of ungerminated seeds with increase in moisture content is almost entirely due to increased respiration of fungi within, or on the surface of, the seed (Crocker and Barton, 1957). 838 FUNGI ATTACKING SEEDS IN DRY SEED-BEDS The failure of the fungicidal dust is in accord with similar experiments with stored grain (Christensen, 1957) and should probably be attributed to the inactivity of the fungicide in the absence of free water. The isolation of P. expansum alone from dusted seeds cannot be interpreted as indicating that this species was the only one active in the seeds. Christensen has shown that although ‘ Ceresan-M’ is not toxic to fungi inside seeds, the residual fungicide on the seed coat, diffusing into the agar, is able to suppress fungal growth from the seed on to agar. to grow out on to agar. Aspergilli and Penicillia isolated from TABLE 5 In the present instance, it is therefore likely that a mixed microflora caused the seed decay but that only P. expansum was able surface-sterilized seeds which had been buried in soil Relative Humidity Week 100 95 90 85 80 1 P. chrysogenum FP. chrysogenum A. glaucus P. crtrinum P. chrysogenum P. citrinum P. citrinum P. chrysogenum P. urticae 2 P. chrysogenum FP. chrysogenum P. chrysogenum P. chrysogenum P. chrysogenum P. urticae P. urticae P. urticae 3 P. expansum P. chrysogenum A. glaucus A. glaucus A. glaucus P. gavanicum P. expansum A. versicolor P. chrysogenum FP. chrysogenum P. urticae P. chrysogenum FP. citrinum P. cttrinum P. citrinum P. rugulosum P. expansum P. nigricans 4 P. expansum P. brevi- A. glaucus A. glaucus A. flavus-oryzae compactum P. janthinellum P. chrysogenum P. chrysogenum P. chrysogenum FP. chrysogenum P. rugulosum P. citrinum P. citrinum P. expansm P. expansum P. nigricans P. expansum P. rugulosum P. janthinellum P. mgricans P. urticae P. rugulosum P. urticae 5 — A. glaucus A. glaucus A. glaucus A. versicolor A. versicolor P. brevi- P. chrysogenum compactum P. chrysogenum P. chrysogenum P. chrysogenum P. citrinum P. citrinum P. expansum P. expansum P. expansum P. expansum P. mgricans P. janthinellum P. javanicum P. javanicum P. rugulosum P. nigricans 7 — P. chrysogenum P. brevi- A. glaucus compactum P. expansum P. chrysogenum P. brev- compactum P. rugulosum P. citrinum P. expansum P. urticae P. janthinellum 9 = A. versicolor A. versicolor A. glaucus A. versicolor P. brevi- P. chrysogenum A. versicolor compactum P. chrysogenum P. expansum P. expansum P. expansum P. javanicum 13 — P. chrysogenum A. glaucus A. glaucus A. glaucus P. expansum P. expansum P, expansum P. brevi- compactum P. rugulosum P. javanicum P. chrysogenum P. urticae P. janthinellum D. M. GRIFFIN 39 Seed decay was rapid in soils at about permanent wilting point or somewhat drier and only in the driest soils did seeds survive for more than three weeks. It seems unlikely that any seed treatment will improve longevity under these conditions, for much of the decay is due to fungi carried inside the testa and at least some of the soil-borne fungi are likely to be tolerant of fungicidal dusts, especially under dry conditions where the compounds are relatively inactive. Acknowledgements I wish to thank Miss E. M. Palmer and Miss C. M. Siddle for laboratory assistance. This work was done whilst I was in receipt of grants from the Wheat Industry Research Council and the N.S.W. Wheat Industry Research Committee. References CHRISTENSEN, C. M., 1957.—Deterioration of stored grains by fungi. Bot. Rev., 23: 108-134. CrocKER, W., and Barton, L. V., 1957.—*~ Physiology of seeds’. Chronica Botanica, Waltham. DommMERGUES, Y., 1962.—Contribution a l’étude de la dynamique microbienne des sols en zone semi-aride et en zone tropicale seche. Ann. Agron. Paris, 13: 265-324, 379-469. GrirFin, D. M., 1963a.—Soil moisture and the ecology of soil fungi. Bzol. Rev., 38: 141-166. , 1963b.—Soil physical factors and the ecology of fungi. III. Activity of fungi in relatively dry soil. Trans. Brit. mycol. Soc., 46: 373-377. Hypz, M. B., 1950.—The subepidermal fungi of cereal grains. I. A survey of the world distribution of fungal mycelium in wheat. Ann. appl. Biol., 37: 179-186. Hyops, M. B., and GatLtEymore, H. B., 1951.—The subepidermal fungi of cereal grains. II. The nature, identity and origin of the mycelium in wheat. Ann. appl. Biol., 38: 348-356. Kovyeas, V., 1964.—An approach to the study of moisture relations of soil fungi. Plant and Soil, 20: 351-363. Rarer, K. B., and Tom, C., 1949.—**‘ A manual of the Penicillia”’. Williams and Wilkins, Baltimore. Snow, D., 1945.—Mould deterioration of feeding stuffs in relation to humidity of storage. Ill. The isolation of a mould species from feeding stuffs stored at different humidities. Ann. appl. Biol., 32: 40-44. Snow, D., Cricuton, M. H. G., and Wricut, N. C., 1944.—Mould deterioration of feeding stuffs in relation to humidity of storage. I. The growth of moulds at low humidities. Ann. appl. Biol., 31: 102-110. STEPHENS, C. G., 1962.—‘‘ A manual of Australian soils”. C.S.I.R.O., Melbourne. Tom, C., and Rarer, K. B., 1945.—“ A manual of the Aspergilli”. Williams and Wilkins, Baltimore. ON THE SPECIES FENESTELLA HOROLOGIA BRETNALL AND MINILYA DUPLARIS CROCKFORD ROBIN WASS Department of Geology and Geophysics, University of Sydney (Plate IT) [Read 27th April, 1966] Synopsis Taxonomic work has established the identity of Fenestella horologia and Minilya duplaris. This identity is discussed and the valid species Fenestella horologia is described. INTRODUCTION During the writer’s work on the Permian Polyzoa of the Bowen Basin, similarities between the species Fenestella horologia and Minilya duplaris became increasingly evident. Both species were described first from the Permian of Western Australia. Fenestella horologia Bretnall, 1926, is characterized by a single row of nodes placed on a carina, hour-glass shaped fenestrules and three zooecia per fenestrule, placed so that there is an aperture at the proximal and distal ends of the fenestrule on the dissepiment and one aperture at mid-length of the fenestrule. Minilya duplaris Crockford, 1944c, the type species of Minilya, is characterized by the same features save for the presence of a double row of nodes on the branch. The nodes arise from a carina which zig-zags its way along the branch. Crockford (1944c, p. 174) notes the difference in carinal appearance, for she states ‘‘ This species (i.e. Minilya duplaris) is probably the same form described by Miss Hosking (1931) from the Wooramel River District as Fenestella horologia Bretnall, but it differs from F. horologia in having a double, instead of a single, row of nodes ”’. The importance of the double row of nodes as a feature of generic significance has been queried by Elias and Condra (1957, p. 66). They produce a very thorough criticism of Crockford’s diagnosis of Minilya and reduce the differences between her genus and Fenestella to one, the double or single row of nodes. Shulga-Nesterenko (1941, pp. 49-51) described Fenestella virgosa Kichwald and its varieties. On F. virgosa var. sparsituberculata only one row of nodes is developed whereas on F’. virgosa a double row of nodes is developed. This variation made the possibility of a single and double row of nodes being developed on the one zoarium appear very real. Recently, Campbell and Engel (1963, p. 67) described Fenestella sp. 1, from the Tournaisian Tuleumba Sandstone, New South Wales, which has both a single and double row of nodes developed. With this in mind and because Hosking (1931) had mistaken Minilya duplaris for Fenestella horologia, measurements of both species were tabulated and plates, as well as primary type material, were studied. The measurements for both species appear below (Table 1). The figures used by Crockford (1944e, p. 181, Pl. 1, figs 6, 7) are reproduced (Pl. IT, figs 1, 2) as they appear in the original plate. From Crockford’s measurements and the figures the similarity between the two species can be observed. PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SoutH WALES, Vol. 91, Part 1 ROBIN WASS 91 _ Additional measurements have been made by the writer on primary types of both species and these are presented in brackets. To determine the base shape of the zooecial chamber, serial sections were cut through Fenestella horologia and Minilya duplaris and camera lucida drawings made. Superposition of the drawings enabled the median plate in the centre of the branch to be related to the nodes on the obverse surface and the change in shape of the zooecial chamber could be studied (see PI. II, figs 3-5). The relation of the nodes to the median plate on Minilya duplaris changes from branch to branch. Because of the similarity between the species, a thorough study was made of all Western Australian material labelled Fenestella horologia or Minilya duplaris. The result was that two specimens were located having the measure- ments tabled and showing both the single and double row of nodes on the one branch. TABLE 1 Fenestella horologia Bretnall Mimlya duplaris Crockford (Crockford, 1944a, 1944c) (Crockford, 1944c) Bw 0-29-0-38 mm. (0-26) 0-33-0-41 mm. B/10 18-22 16-19 (21) F/10 16-18 14-17 (18) Fl 0-29-0-52 mm. 0-4-0-51 mm. Fw 0-17-0-4 mm. 0-14-0-25 (0-32) mm. Z/10 37 (35-39) 33 (37) Zd 0-08—0-13 mm. 0-13 mm. Z/E 2-3 2-3 Z-Z 0:21-0:34 mm. 0-27-0-35 mm. N-N 0:23-0:31 mm. 0-13-0-17 mm. N/5 (16-21) i (30-38) Dw 0-1—0-29 mm. 0-1—0-3 mm. Zb (4) (T) Bw=branch width, B/10=branches per 10 mm., F/10=fenestrules per 10 mm., Fl=fenestrule length, Fw—fenestrule width, Z/10—zooecia per 10 mm., Zd=zooecial diameter, Z/F=zooecia per fenestrule, Z-Z= separation of centres of successive zooecial apertures, N-N=separation of centres of successive nodes, N/5=nodes per 5 mm., Dw=dissepiment width, Zb—zooecial base shape, T=trapezoidal. One specimen, F.17538 Australian Museum Collection, is labelled Fenestella horologia but for the most part it shows the double row of nodes of Minilya duplaris. Measurements made on the zoarium showing the double row of nodes agree with the measurements tabled previously (see Pl. IJ, fig. 6). Of interest was the spacing of the nodes. On the portion of the zoarium where a double row of nodes is a constant feature, the spacing varied from 0:12 mm. to 0-17 mm. The spacing in the first appearance of a single row varied from 0:14 mm. to 0-17 mm. This was not into the spacing of Fenestella horologia but unfortunately the single row condition does not persist for more than three nodes before reverting to a double row condition and then to a single row condition of similar length with the nodes separated by 0-14 mm. to 0-2 mm. Another observation which can be made is that the two cases of the single row have the same orientation with respect to a straight line, even though they are separated by an area showing the double row condition. A second specimen, labelled C.P.C. 1287c, was assigned to Minilya duplaris by Crockford (1957, p. 67). This specimen likewise has both a single and double row of nodes developed (see Pl. II, fig. 7). 92 THE SPECIES FENESTELLA HOROLOGIA AND MINILYA DUPLARIS Clearly the two species are conspecific and therefore, because of priority, Minilya duplaris is invalid. As this is the type species of the genus Minilya, the genus is also invalid. There are not many specimens of Minilya amplia Crockford, 1944), available for study but it is felt that when additional specimens are found this species will be identical with a species of Fenestella having both a single and double row of nodes developed. SYSTEMATIC DESCRIPTION Order CRYPTOSTOMATA Shrubsole and Vine, 1882 Family FENESTELLIDAE King, 1850 Genus FENESTELLA Lonsdale, 1839 Type Species.—(by subsequent designation of Riley, 1962, p. 76) Fenestella subantiqua WOrbigny, 1850, p. 180, from the Silurian, Wenlockian, Wenlock Limestone, at Wren’s Nest, Dudley, Worcestershire, England. Diagnosis.—Zoarium fan or funnel shaped; zooecia in two rows on the branches, commonly increasing to three, proximal to bifurcation; rows of zooecia separated by a nodose carina on the obverse surface; reverse surface of varying ornamentation. FENESTELLA HOROLOGIA Bretnall, 1926 1926, Fenestella horologia Bretnall, p. 15, Pl. 1, fig. 6. 1929, Fenestella parviuscula Bassler, p. 76, Pl. cexli, figs 8-13. 1931, ? Fenestella parviuscula Bassler; Martin, p. 391. 1931, Fenestella horologia Bretnall; Hosking, p. 13, Pl. 4, fig. 3. 1932, Fenestella parviuscula Bassler; Fritz, p. 99. 1936, Fenestella pectinis Moore (partim); Chapman; in Raggatt, p. 128 (fide Crockford, 1944¢, p. 158). 1937, Fenestrellina parviuscula (Bassler); Elias, p. 314. 1943, Fenestrellina horologia (Bretnall) ; Crockford, p. 266. 1944a, Fenestrellina horologia (Bretnall) ; Crockford, p. 189, Pl. 1, fig. 1; Pl. Zhe AC 1944b, Fenestrellina horologia (Bretnall) ; Crockford, p. 158. 1944¢, Fenestrellina horologia (Bretnall); Crockford, p. 167, Pl. 1, figs 3, 6. 1944¢c, Minilya duplaris Crockford, p. 173, Pl. 1, figs 5, 7; text-figs 1C, D. 1946, Minilya duplaris Crockford ; Crockford, p. 132. 1951, cf. Fenestella ivanovi Shulga-Nesterenko, p. 100, Pl. 19, fig. 1; text- fig. 38. 1953, Fenestrellina nodograciosa Chronic; Chronic, in Newell, Chronic and Roberts, p. 111, Pl. 21, figs 4a, b. 1955, cf. Fenestella donensis Morozova, p. 27, Pl. 4, figs 2, 3; text-figs 5, 6. 1957, Fenestella horologia Bretnall; Crockford, p. 57. 1957, Minilya duplaris Crockford ; Crockford, p. 67. 1957, Minilia duplaris [sic] Crockford ; Elias and Condra, p. 65. 1957, Fenestella parviuscula Bassler; Elias and Condra, p. 108. 1961, Fenestella nomatae Sakagami, p. 34, Pl. 15, fig. 3; text-fig. 5. Holotype.—Specimen 16, Western Australian Geological Survey Collection from the Gascoyne River District, Western Australia. The label was lost; there are a number of specimens of the species on the block figured by Bretnall (1926, Pl. 3) (Crockford, 1944a, p. 190). Neotype.—(chosen Crockford, 1944a, p. 190) No. 2/2405¢e, Western Australian Geological Survey Collection from the Permian, Sakmarian, Callytharra Formation, east of the Gascoyne-Wyndham River Junction, Western Australia. ROBIN WASS 93 Lat. 25°03’, Long. 115°33’, Glenburgh 1 : 250,000 Geological Series Sheet SG 50-6, at Fossil Hill, Fossil Hill Station, Western Australia (Crockford, 1944a, p. 190, Pl. 1, fig. 1). This specimen is presently housed at the Depart- ment of Geology and Geophysics, University of Sydney. Diagnosis.—Species of Fenestella with hour-glass fenestrules, two or three zooecia per fenestrule ; the relationship between zooecia and dissepiments is stabilized ; nodes may be uniserial or biserial ; zooecial base shape trapezoidal. Discussion.—Crockford (1944a, p. 190) states that the specimen chosen as neotype is part of a block from the Gascoyne River District which was used by Bretnall for his original description. The specimen is designated ‘‘ neotype ” because there is doubt whether it was a syntype. The specimen chosen and figured by Crockford as neotype can be seen on one of Bretnall’s plates (1926, Pl. 3). In the lower half of this plate, there is a figure ‘‘1”’ on its side and the neotype is half an inch from the lower right-hand corner of this figure. As recently as 1964, Crespin (1964) quotes the type locality of the neotype as being between the top of the Lyons Series and the top of the Byro Series, Gascoyne River District, Carnarvon Basin, Western Australia. The stratigraphic thickness represented is of the order of 3,500 feet. Because of the uncertainty, an effort was made to obtain more information concerning the type locality and to quote it as accurately as possible. The locality given for the neotype is the result. With reference to the Glenburgh 1 : 250,000 Sheet, some mention must be made regarding the type locality and the stratigraphic horizon. Although the Callytharra Formation is not shown as outcrop at Fossil Hill, M. A. Condon (pers. comm.) has assured the writer that the Formation is present at this locality. Description. Micrometrie Formula B/10 D/10 Z/5 N/5 Bw Zd Zb 18-22 15-18 17-20 17-21 0-3-0 -38 0 -08-0 -13 T The branches are not parallel and show a tendency to diverge. They are capped by a nodose carina. Proximal to bifurcation, the branch width may increase to 0-48 mm.; distal to bifurcation, the branch assumes the usual proportions. The branches are of similar width to the fenestrules. The fenestrules have a width of 0-17 mm. to 0-40 mm. and a length of 0:29 mm. to 0:52 mm. The fenestrules are hour-glass in shape because of the projection of the peristomes of the apertures into them. The dissepiments vary in width from 0-10 mm. to 0-30mm. Their width increases near the branch because an aperture is situated on the dissepiment. The dissepiments are perpendicular to the branches. The zooecial apertures are in two rows, with the zooecia of one row alternating with those of the other. There are three zooecial apertures per fenestrule and in many cases two of them are partly or wholly situated on a dissepiment. The centres of the zooecial apertures are separated by 0-21 mm. to 0-34 mm. Well developed peristomes are evident on the apertures, more so on the side adjacent to the fenestrule than on the side adjacent to the carina. Proximal to bifurcation, a third row of apertures is inserted but no case has been found of four rows of apertures. The third row of apertures is located distal to the bifurcation of the carina. The carina is nodose and has one row of nodes developed which exhibit varying tendencies. They may be bifid or even trifid at their topmost extremity or they may not bifureate. The centres of successive nodes are 0-23 mm. to 0-31 mm. apart. 94 THE SPECIES FENESTELLA HOROLOGIA AND MINILYA DUPLARIS The zooecial base shape in the upper portions of the chamber is triangular but at the base it is trapezoidal. On the reverse surface both branches and dissepiments are rounded. Ornamentation of the outer layer consists of minute, randomly oriented nodes. The removal of this layer reveals longitudinal striations. Remarks.—Species with similar meshwork formulae to f. ire are F. subquadratopora Shulga-Nesterenko, 1952, and Ff. subvischerensis Shulga- Nesterenko, 1951. The former species has a different arrangement and spacing of the nodes, while the latter species has narrower branches and dissepiments and the hour-glass fenestrules are not well developed. Fenestella girtyi (Elias), 1937, has similar dimensions to F. horologia but there is a difference in the base shape of the zooecial chamber and the relation between the apertures and the dissepiments is not well stabilized in F. girtys whereas it is in F. horologia. Fenestella vischerensis Nikiforova, 1938 (fide Shulga-Nesterenko, 1941), and F. vischerensis var. baschkirica Shulga-Nesterenko, 1941, have narrower dissepi- ments and branches, the latter feature being characteristic also of F. micro- aperturata Shulga-Nesterenko, 1941, and F. microaperturata var. polaris Shulga- Nesterenko, 1941. F. microaperturata also has a smaller zooecial diameter. Range and Distribution.—Fenestella horologia Bretnall has been recorded previously from Western Australia (Bretnall, 1926 ; Crockford, 1951 ; Crockford, 1957). It ranges from the Callytharra Formation of Sakmarian age to the Liveringa Formation of Kungurian age. As F. parviuscula, it has been recorded from the Bitaoeni and Basleo Beds of Timor. These are Artinskian-Kazanian in age; as /’. nodograciosa (Chronic), it has been recorded from the Upper Pennsylvanian of Peru, and as F. nomatae Sakagami, it occurs in the Upper Permian of Japan. From the Artinskian of Vancouver Island, Canada, it is listed as F. parviuscula Bassler by Fritz. In Eastern Australia, Crockford (1943, 1946) has recorded the species from the Lake’s Creek Quarry, east of Rockhampton, and from Consuelo Creek, south-west of Rolleston. During work by the writer on the Permian Polyzoa of the Bowen Basin, it has been recorded from the Artinskian Buffel Formation, south of Cracow, from the Yatton Limestone, north-west of Marlborough, and from the Artinskian Cattle Creek Formation in the Reid’s Dome area, south- west of Rolleston. Acknowledgements The writer would like to thank Dr. T. B. H. Jenkins for discussion on the subject and for critical reading of the manuscript, and Dr. J. M. Dickens for discussion on the type locality. References BaASSLER, R. 8., 1929——The Permian Bryozoa of Timor. Paldont. Timor, 16: 37-90. BRETNALL, R. W., 1926.—Palaeontological contributions to the Geology of Western Australia, 7 (13): Descriptions of some Western Australian Fossil Polyzoa. Bull. geol. Surv. West. Aust., 88: 7-33: CAMPBELL, K. 8S. W., and EnceL, B. A., 1963.—The faunas of the Tournaisian Tuleumba Sandstone and its members in the Werrie and Belvue Synclines, New South Wales. J. geol. Soc. Aust., 10: 55-122. Crespin, I., 1964.—Catalogue of type and figured specimens in Western Australia. Rep. Bur. Miner. Resour. Geol. Geophys. Aust., 71. CRocKFORD, J. M., i yozoa from Hastern Australia, Part II]: Batostomellide and Fenestrellindae from Queensland, New South Wales and Tasmania. J. Proc. r. Soc. N.S.W., 76: 258-267. , 1944a.—A revision of some previously described species of Bryozoa from the Upper Palaeozoic of Western Australia. J. Proc. R. Soc. West. Aust., 28: 187-199. ROBIN WASS 95 CrocKkForD, J. M., 1944b.—Bryozoa from the Permian of Western Australia, Part I: Cyclostomata and Cryptostomata from the North-West Basin and Kimberley District. Proc. Linn. Soc. N.S.W., 69: 139-175. , 1944c.—Bryozoa from the Wandagee and Noonkanbah Series (Permian) of Western Australia. J. Proc. R. Soc. West. Aust., 28: 165-185. , 1946.—A bryozoan fauna from the Lake’s Creek Quarry, Rockhampton, Queensland. Proc. Linn. Soc. N.S.W., 70: 125-134. —, 1951.—The development of bryozoan faunas in the Upper Palaeozoic of Australia. Ibid., 76: 105-122. , 1957.—Permian Bryozoa from the Fitzroy Basin, Western Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., 34. Eras, M. K., 1937.—Stratigraphic significance of some Late Palaeozoic Fenestrate Bryozoans. J. Paleont., 11: 306-334. , and Conpra, G. E., 1957.—Fenestella in the Permian of West Texas. Mem. geol. Soc. Am., 70. Fritz, M. A., 1932.—Permian Bryozoa from Vancouver Island. Trans. r. Soc. Can., Ser. 3, 26: 93-110. Hosxine, L. F. V., 1931.—Fossils from the Wooramel District, Western Australia. J. Proc. r. Soc. West. Aust., 27: 7T-5d2. ; Marttn, R., 1931.—De Palaeontologie en Stratigraphie van Nederlandsch Oost-Indie: Bryozoa. Leid. geol. Meded., 5: 390-395. Morozova, I. P., 1955.—Carboniferous Bryozoa of the Middle Don Basin. Trudy paleont. Inst., 58. NeEweE Lt, N. D., CuHronic, J., and Roperts, T., 1953.—Upper Palaeozoic of Peru. Mem. geol. Soc. Am., 58. Raceatt, H. G., 1936.—Geology of the North-West Basin, Western Australia, with particular reference to the stratigraphy of the Permo-Carboniferous. J. Proc. r. Soc. N.S.W., 70: 100-174. Riney, N. D., 1962.—Opinion 622: Fenestella Lonsdale, 1839 (Class Bryozoa); Validation under the Plenary Powers in accordance with the accustomed usage. Bull. zool. Nom.., 19: 76-79. SaKkaGami, S., 1961.—Japanese Permian Bryozoa. Spec. Pap. Palaeont. Soc. Japan, 7. SHuLGA-NESTERENKO, M. I., 1941.—Lower Permian Bryozoa of the Urals. Paleontologiya SSSR, 5 (5): 1. ————, 1951.—Carboniferous Fenestellids from the Russian Platform. Trudy paleont. Inst., 32. , 1952.—New Permian Bryozoa from the Uralian Region. JIbid., 37. , 1955.—Carboniferous Bryozoa of the Russian Platform. Jbid., 57. EXPLANATION OF PLATE II Fig. 1. Fenestella horologia Bretnall ; reproduced from Crockford, 1944c, Pl. 1, fig. 6. (nodes retouched). x 20. Fig. 2. Minilya duplaris Crockford ; holotype, reproduced from Crockford, 1944c, Pl. 1, fig. 7. (modes retouched). x 20. Fig. 3. Camera lucida drawing of Fenestella horologia Bretnall to show relation of nodes, median plate and apertures. x 25. Fig. 4. Fenestella horologia Bretnall ; camera lucida drawing to show the shape of the zooecial base chamber and relation to apertures. x 25. Fig. 5. Camera lucida drawing of Minilya duplaris Crockford to show the relation of nodes, median plate and apertures. x 25. Fig. 6. Fenestella horologia Bretnall; specimen F.17538, Aust. Mus. Collection, showing uniserial and biserial arrangement of nodes (retouched). x 25. Fig. 7. Minilya duplaris Crockford; specimen C.P.C. 1287¢ showing uniserial and biserial arrangement of nodes (modes retouched). x 20. TWO NEW SPECIES OF PERMIAN BRACHIOPODS FROM QUEENSLAND ROBIN WASS Department of Geology and Geophysics, University of Sydney (Plate III) [Read 27th April, 1966] Synopsis Cancrinella gyrandensis, sp. nov., and Grantonia cracovensis, sp. nov., are described from Permian strata at Cracow, Queensland. INTRODUCTION During a study of Permian sediments and faunas in the Cracow District in the south-eastern part of the Bowen Basin, Queensland, Cancrinella gyrandensis and Grantonia cracovensis were identified from the Barfield and Buffel Formations respectively. The species of Cancrinella, C. gyrandensis, is regarded as being identical with the form figured by Campbell (1953) as Canerinella cf. magniplica sp. nov. from a locality north of Cracow. This work is based on portion of a thesis submitted as partial fulfilment for the degree of B.Sc. with Honours in Geology of the University of Queensland. All fossil specimen numbers prefixed by “F” and locality numbers prefixed by “L” belong to the University of Queensland, Department of Geology Catalogues. Map references refer to the Mundubbera 1: 253,440 Military Map. SYSTEMATIC DESCRIPTIONS Phylum BRACHIOPODA Class ARTICULATA Suborder PRODUCTOIDEA Mailleux, 1940 Superfamily PRODUCTACEA Waagen, 1883 Family LINOPRODUCTIDAE Stehli, 1954 Subfamily LINOPRODUCTINAE Stehli, 1954 Genus CANCRINELLA Fredericks, 1928 1928, Cancrinella Fredericks, pp. 784, 791. 1932, Canerinella Dunbar and Condra, p. 257. 1937, Cancrinella Sarytcheva, pp. 78, 110. 1950, Cancrinella Hill, p. 12. Type Species—(By original designation) Productus cancrini de Koninck, 1842, p. 179, Pl. 9, figs 3a, b, from the Zechstein strata on the banks of the River Kidash, near Bielibei, Orenburg, Russia. Diagnosis.— Small, thin shelled productids with ventral valve convex and without sinus, and dorsal valve concave or slightly geniculated. Ornament of thin, radial striae and concentric wrinkles, and spines, scattered on the ventral area and always along the hinge line and on the ears. Hinge line without area or teeth, with cardinal process in the form of two loops on an elevated ridge in the dorsal valve. Median septum well defined.” (After Sarytcheva, 1937). Discussion.—For discussion of the type species, the reader is referred to Hill (1950), Bryant (1955) and Waterhouse (1964). PROCEEDINGS OF THE LINNEAN SOCIETY OF NEW SoutTH WatLEs, Vol. 91, Part 1 ROBIN WASS 97 CANCRINELLA GYRANDENSIS, sp. Nov. (Pl. ITI, figs 1-5) 1953, Cancrinella cf. magniplica Campbell, p. 29. Holotype.—¥.43422 from the Barfield Formation, north-west of Cracow, Queensland; 1.2573, 31628589, Par. Coteeda, Co. Dawson, 0-7 mile along the road to ‘‘ Gyranda’”’ from the Theodore Road, north of Cracow. Paratypes.—¥.43423, F.43424 from the above locality. Diagnosis.—Very large Cancrinella with large visceral cavity and strong transverse wrinkling on the trail. Description.—The shell is concavo-convex, of large size and elongate. The visceral cavity is large. Hinge line is straight and in length is generally a little less than the maximum width of the shell. The trail is long. The ventral valve is very convex, the greatest convexity being at the umbo. The valve is more convex in transverse than in longitudinal profile. The umbo is incurved over the hinge line and the umbonal slopes are steep. The ears are small, slightly produced and flat. The venter is non-sinuate and gently rounded or flattened. The ornament consists of fine radial capillae which increase by intercalation. They number from 28 to 33 in 10 mm. on the trail. The entire ventral surface is covered by transverse wrinkles, which are more strongly developed on the trail. Here they are up to 1-5 mm. high and extend across the valve. Spines are developed by an increase in the anterior end of a short raised portion of the capillae. Because of the nature of the surface, the arrangement of the spines cannot be determined. The dorsal valve is concave, the region of greatest concavity being at the anterior end of the visceral disc. The curvature is approximately the same as in the ventral valve. The valve is sharply geniculate. The ears are flattened and differentiated from the visceral disc by a slight change in slope at 30° to the hinge line. The ornament of the dorsal valve is similar to that of the ventral valve except that the transverse wrinkling is not as strongly developed. The internals of both valves are poorly known. The visceral disc is two-thirds as wide as it is long. It does not vary greatly in proportion. A low Median septum is developed in the dorsal valve. Remarks.—This species has affinities with C. magniplica and is the same as those specimens figured by Campbell (1953, p. 29, Pl. 1, figs 6-8) as C. ef. magniplica from an erosional gully, halfway between Oxtrack and Delusion Creeks on the Cracow-Theodore Road. C. magniplica has a much smaller visceral cavity and weaker wrinkles on the trail. It is more finely costate. C. farleyensis (Etheridge and Dun), 1909, is more coarsely costate, with a smaller visceral cavity. It is less strongly wrinkled. Both the previously mentioned species are smaller in size than C. gyrandensis. Campbell (1953) stated that a trend to coarser wrinkling was initiated in the Cattle Creek Formation and carried into the Ingelara Formation with accompanying decrease in the size of the costae. This is not the case as C. gyrandensis is more strongly wrinkled and has coarser costae than C. magniplica, even though it is found at a higher stratigraphic horizon than C. magniplica. However, a trend towards coarser transverse wrinkling may be associated with a tendency to increase the size of the visceral cavity as higher stratigraphic horizons are reached. Of the overseas species, C. rugosa Cloud, 1944, from the Mid Guadalupian of U.S.A., shows some resemblance to this species. It is of slightly smaller size and does not have the strong transverse wrinkling. It is more coarsely costate. 98 TWO NEW SPECIES OF PERMIAN BRACHIOPODS FROM QUEENSLAND Range and Distribution.—The writer has found the species only in the Barfield Formation in the Theodore-Cracow District. J. M. Dickins (pers. comm.) has advised the writer that he has not found the species elsewhere in the Bowen Basin. Localities.—In addition to the previously mentioned localities, C. gyrandensis is recorded from the Barfield Formation at L.2572, 31698408, Par. Cracow, Co. Dawson, approximately 4-0 miles south-west of ‘*‘ Cracow”. Specimens from this locality are F.43577-43581. Dimensions.—(Ventral valve) F.43424 F..43423 F..43422 Length 67+ 68 + (sya Width (Maximum) 38 40 37 mm. Suborder SPIRIFEROIDEA Allen, 1940 Superfamily SPIRIFERACEA Waagen, 1883 Family SPIRIFERIDAE King, 1846 Subfamily TRIGONOTRETINAE Schuchert, 1898 Genus GRANTONIA Brown, 1953 1953, Grantonia Brown, p. 60, Pl. vi, figs 1-8. Type Species—(By original designation) Grantonia hobartensis Brown, 1953, Pl. vi, figs 1-8, from the Permian, Artinskian, Berriedale Limestone, in Rathbone’s Quarry, New Norfolk Road, near Granton, about 12 miles north-north-west of Hobart, Tasmania. Diagnosis.—See Brown, 1953, p. 60. Discussion.—The genus Grantonia was erected by Brown for specimens of Trigonotreta stokest auctt. (non Koenig, 1825). Specimens referred to Spirifer tasmaniensis Morris, 1845 are synonyms of Trigonotreta stokesi Koenig, 1825. Differences between Grantonia, Neospirifer Fredericks, 1924, Trigonotreta Koenig, 1825, and Spirifer Sowerby, 1818, may be seen in ornamentation and gross form. The writer (Wass, 1965) has discussed the differences between the first three genera. Range and Distribution.—In Tasmania, the genus is found in the Quamby, Golden Valley and Cascades Groups and the Malbina Formation of Lower Permian age. In Queensland, the writer has found the genus only in sediments of similar age which are equivalent to the Cattle Creek Formation. GRANTONIA CRACOVENSIS, Sp. nov. (Pl. ITI, figs 6-11) Holotype.—F¥.43392, from the Buffel Formation, Cracow, Queensland ; L.2575, 32128451, Par. Cracow, Co. Dawson, at the base of the ridge, half a mile north-west of ‘‘ Cracow ”’, six miles south of Cracow, Queensland. Paratypes.—¥.43393, F.43396—43398 from the above locality. Diagnosis.—Species of Grantonia with four plicae developed on each side of the sulcus and nine costae in the suleus; crural lamellae divergent. Description.—The valves are biconvex and semi-circular in outline; the maximum width is at the hinge line. In the ventral valve, the greatest convexity is at the umbo where it overhangs the hinge line. The ventral sulcus is evident at the umbo and widens greatly anteriorly with an associated increase in depth anterior to the muscle platform. The cardinal extremities are almost rounded; the cardinal areas are broad, triangular and striated both longitudinally and transversely, with the former being more pronounced. The areas are convex to the anterior margin. ROBIN WASS 99 The fine costae are spread over four plicae on each side of the sulcus. The ventral sulcus has a median costa with four costae of similar size developed on each side. Hach side of the sulcus is bordered by a raised costa of larger size. On the plicae adjacent to the sulcus, bifurcation begins to take place 8 mm. from the umbo, but on the more lateral plicae bifurcation is absent until 13 mm. from the umbo. The entire surface is ornamented by concentric growth lines. The high dorsal fold is bounded on both sides by four plicae and the surface is multicostate; on each side of the fold there are four costae but this may be increased to five anteriorly. Due to the slightly distorted nature of the external ornament, it is difficult to state accurately the number of costae on each side of a plication. In the anterior region there may be three. The external ornamentation is reproduced on an internal mould. In the ventral valve there is a wide, open delthyrium bounded on each side by dental lamellae which converge towards the centre of the valve and do not reach the floor. At their extremities they produce small, pyramidal projections which fit into sockets in the dorsal valve; the projections taper off very sharply anteriorly. Heavy callus is present in the delthyrium and in the umbonal region ; this callus is differentiated from the cardinal area by two prominent ridges that diverge at an angle of 45° from the umbo. The long, narrow adductor scars are raised above the heart-shaped diductors which border them. The adductors are striated longitudinally ; on the diductors, the striations are mainly concave posteriorly but in the anterior portion of the scar they tend to become convex posteriorly. In the dorsal valve, the cardinal process is composed of numerous fine, parallel plates which parallel the median plane of the valve. Flanking the cardinal process are dental sockets which taper posteriorly and make an angle of 90° with one another. The dental sockets border two thin lamellae which diverge towards one another anteriorly and give rise to the crural plates. They do not seem to reach the floor of the valve. A median septum extends half the length of the dorsal valve. Remarks.—Grantonia cracovensis is easily distinguished from the type Species by the diverging crural lamellae, the ornament and a wider muscle field in the former species. Specimens from Beds 8 and 9 at Homevale (Mt. Britton) have been examined and they are considered to be identical with G. cracovensis. Preservation of the forms described by Maxwell (1964) as Grantonia cf. hobartensis is too poor to allow definite comparisons to be made with the Cracow material. Range and Distribution.—The species has been found only in Queensland at Cracow and Mt. Britton in strata of Artinskian age. Localities.—In addition to the previously mentioned locality, G. cracovensis is found in the Buffel Formation at 1.2483, 3175 8603, 0-8 mile north-west of Rose’s Pride Mine. The specimens from this locality are F.43595—43597. Dimensions.—(Ventral valve). F.43392 F.43393 F.43398 Length 36 39 36 mm. Width 52 59 54 mm. Height 13 16 14 mm. 100 TWO NEW SPECIES OF PERMIAN BRACHIOPODS FROM QUEENSLAND Acknowledgements The writer wishes to thank Professor Dorothy Hill, F.R.S., of the University of Queensland, who supervised the project, and Dr. T. B. H. Jenkins of the University of Sydney for his critical reading of the manuscript. Photographie work is by the Photography Department, University of Queensland. References Brown, I. A., 1953.—Permian Spirifers from Tasmania. J. Proc. r. Soc. N.S.W., 86: 55-63. Bryant, D. L., 1955.—Index, type species and bibliography of Productid genera. J. Paleont., 29: 283-294. CAMPBELL, K. 8. W., 1953.—The fauna of the Permo-Carboniferous Ingelara Beds of Queensland. Pap. Dep. Geol. Univ. Q’land., 4 (3). CLoup, P. E., in Kine, R. H., DunBAR, C. O., CLoup, P. E., and Mituer, A. K., 1944.— Geology and Palaeontology of the Permian area, north-west of Las Delicias, South-Western Coahuila, Mexico. Spec. Pap. geol. Soc. Am., 52: 49-69. Dunpar, C. O., and Conpra, G. EH., 1932——Brachiopoda of the Pennsylvanian System in Nebraska. Bull. Neb. geol. Surv., 2. ETHERIDGE, R., and Dun, W. S8., 1909.—Notes on the Permo-Carboniferous Producti of Hastern Australia. Rec. geol. Surv. N.S.W., 8: 293-304. FREDERICKS, G., 1928.—Contribution to the classification of the genus Productus. Bull.. geol. Com., 46: 773-792. Hix, D., 1950.—The Productinae of the Artinskian Cracow fauna of Queensland. Pap. Dep. Geol. Univ. Q’land., 3 (2). DE Konincx, L. G., 1842.—Description des animaux fossiles qui se trouvent dans le terrain carbonifere de Belgique. Liége. Maxwetit, W. G. H., 1964—The geology of the Yarrol region. Part 1: Biostratigraphy. Pap. Dep. Geol. Univ. Q’land., 5 (9). SaRyTcHEVA, T. G., 1937.—Lower Carboniferous Producti of the Moscow Basin. Trav. Inst. paleozool. Acad. Sci. U.R.SS., 6. Sten, F. G., 1954.—Lower Leonardian Brachiopoda of the Sierra Diablo. Bull. Am. Mus. nat. Hist., 105: 365-385. Wass, R. E., 1966.—The marine Permian Formations of the Cracow District, Queensland. J. Proc. r. Soc. N.S.W., 98, Part 3: 159-167. WatERHOUSE, J. B., 1964.—Permian Brachiopods of New Zealand. Palaeont. Bull., Wellington, 35. EXPLANATION OF PLATE III Figs 1-5. Cancrinella gyrandensis sp. nov. 1, F. 43422 from L. 2573. x1; 2, F. 43422 from L. 2573. x1; 3, F. 43422 from L. 2573. x1; 4, longitudinal section of F. 43424 from L. 2573. x1; 5, transverse section of F. 43423 from L. 2573. x1. Figs 6-11. Grantonia cracovensis sp. nov. 6, F. 43398 from L. 2575. x1; 7, F. 43392 from LL. 2575. x1; 8, internal of ventral valve of F. 43393 from L. 2575. x1; 9, ornament of ventral valve of F. 43396 from L. 2575. x1; 10, internal of dorsal valve of F. 43397 from L. 2575. x1; 11, internal of dorsal valve of F. 43397 from L. 2575. x1. Proc. Linn. Soc. N.S.W., Vol. 91, Part 1 PLATE I kc/sec seconds Fig. 1. Audiospectrograms of part of call of A, Hyla jervisiensis; and B, H. ewingi. Fig. 2 Adult male Hyla jervisiensis. ie ~? a Mle Proc. Linn. Soc. N.S.W., Vol. 91, Part 1 PLATE II Henestella horologia and Minilya duplaris, ROC MINN SOC. IN:S We Viol Qik Part 1 PLATE I Canerinella gyrandensis and Grantonia cracovensis, SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1966 THE CENTENARY OF MENDEL D. G. CATCHESIDE John Curtin School of Medical Research, Australian National University, Canberra [Delivered 15th July, 1966] Mendel, the centenary of whose work is the subject of this Sir William Macleay Memorial Lecture, had the unusual distinction of originating a new branch of science. Generally, in the development of science, a new advance depends on many predecessors whose work provides a basis for it. Genetics is an exception, owing its origin to one man, Gregor Johann Mendel, who expounded its principles at Brno in two lectures on 8th February and 8th March, 1865. His work made scarcely a ripple in the tumultuous sea of biological study, so stirred by the tidal waves of Darwinism in 1859. It was forgotten for the next 35 years, barely even noticed and certainly not understood. Mendel had no authority as a biologist and the very merits of his work, as we see it, must have made it suspect at the time and seemingly the work of an eccentric. Even the very simplest mathematical practices, beyond mere measurement, had found no place in biology. A predecessor is a man who, at an earlier date, makes a discovery which his successor is able to enlarge into a general principle of universal validity. In this sense Mendel had no predecessors. There were many breeders who hybridised different species and varieties of plants and domesticated animals. However, the objectives they were studying do not really constitute basic genetics and claims that any of them had anticipated Mendel do not bear inspection. They were studying hybridism, whether sterility in a hybrid was due to the pollen parent or to the seed parent, whether either parent could be held responsible for the characters of different parts of the plant, which parent had prepotency in determining the characters of the hybrid, whether the parents were different species or merely varieties. Thus Colladon in 1822 was led to conclude that grey and white mice were different species! Generally, the parents used by the early hybridisers in their experiments were either different species or else varieties differing in many characters. The results obtained were chaotic and often inconstant or contradictory. They led to no general principles. This was the difference between previous attempts to study heredity and the revolution effected by Mendel. The failure of his contemporaries to recognise the significance of Mendel’s work confirms that there were no minds already prepared to see that the problem of heredity had been laid bare to complete solution. None saw that Mendel’s researches supplied what he, rightly, considered to be missing from Darwin’s theories, namely a mechanism of heredity and of the conservation of heritable variation. Indeed, the influence of Nageli, interested in the same trivialities as the early hybridisers, led Mendel to fruitless and exacting work in hybridising hawkweeds, now known to lack sexual reproduction. No earlier work anticipates Mendel’s reduction of the problem of heredity to the study of single character differences which result, after crossing, in hybrid offspring in certain specified proportions. Goss, indeed, working also with garden peas, had observed dominance, recessiveness and segregation, going so far as to count the numbers of the different kinds in the F,. Likewise, Darwin observed these features in snapdragons. Neither arrived at any general principles as a consequence. PROCEEDINGS OF THE LINNEAN SociETy oF NEw SoutH Watss, Vol. 91, Part 2 “= 102 SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1966 Mendel, in writing about previous work, clearly stated his new scientific procedures as follows: ‘“‘. .. among all the numerous experiments made, not one has been carried out to such an extent and in such a way as to make it possible to determine the number of different forms under which the offspring of hybrids appear, or to arrange these forms with certainty according to their Separate generations, or definitely to ascertain their statistical relations ”’. The constancy of characters of hybrids, as Mendel explained, was the starting point of his work. It enabled him to postulate an explanation of his results by supposing that the characters were each determined by particulate factors, that the factors from each parent segregated completely at the formation of the germ cells and in equal numbers, and that there were equal chances of fertilisation of germ cells bearing either one of the two different factors, which thereby became recombined. In fact, Mendel made scarcely any distinction between the unit characters visible in the adult organisms and the unit factors borne in their germ cells, using the word Merkmal for both and treating them as a virtual identity. He left out everything that comes between them, how one determines the other, the whole of development. Knowledge of the connections is immaterial to an understanding of heredity. Statistical analysis of Mendel’s data led Fisher, in 1930, to conclude that Mendel knew what proportions to expect when he made his experiments. This means that he had thought out his scheme of particulate inheritance beforehand, as an abstract and simple combinatorial exercise, before putting it to the test. It was not an empirical discovery subsequently processed by induction into a scheme. Curiously, in reporting his work to the Natural History Society in Brno, Mendel presented the data at the first lecture and the mathematical explanation at the second one. The statistical evidence is that Mendel’s data agree much better with expectation than can reasonably be expected. Taken together, the figures published by Mendel would be expected to be equalled or improved upon, in their agreement with expectation, only once in 8,000 trials. There is no question of attributing deliberate falsification to Mendel or to his assistants, to whom Mendel’s expectations would have been known. But enthusiastic amateurs are prone to give an experiment the benefit of a doubt, and to bias results unintentionally by discarding poor specimens and doubtful cases, to count a portion of a sample, stopping when the proportions are ‘‘ good ’’, thus making honest mistakes. Perhaps, indeed, Mendel did observe just this unusually consistent set of data. The conclusion that Mendel’s work stemmed from an idea, worked out theoretically as a mathematical exercise, is supported by the very thorough manner in which Mendel expressed his system in mathematical terms and in a general form for universal application. All of the main features of Mendel’s theory would follow from the simplest possible assumptions of particulate inheritance. To conceive this simple hypothesis is exactly what Mendel, failed twice in an examination to qualify as a high school teacher, appears to have done. Moreover, the experiments to test the theory were performed with the most meagre facilities, a garden 35 x7 metres, a few varieties of the garden pea (Pisum sativum), some forceps and bags to exclude insects from the flowers. Mendel’s theory could hardly have been formulated without some knowledge of the facts of fertilisation, that one egg and one sperm unite to form the zygote. This became known only a few years before his work and was not generally accepted even then. Thus Darwin thought more than one sperm was needed for each egg, both in animals and plants. Direct observations of fertilisation were dependent upon the development of compound microscopes. Amongst lower plants fertilisation was observed in Fucus by Thuret in 1853, in Oedogonium by Pringsheim in 1856 and in fungi by De Bary in 1861. The role of the nucleus in fertilisation and cell division D. G. CATCHESIDE 103 was established by the work of Hertwig in 1875 on the fertilisation of the egg of the sea urchin. In seed plants, Amici in 1823 observed the production of the pollen tube, which he traced in 1830 to the ovary and even to the micropyle. In 1846 he showed, in orchids, that a cell, already present in the ovule and inactive until the pollen tube arrives, then develops into the embryo. This was confirmed and extended by Hofmeister. Hence Mendel could state ‘ In the opinion of renowned physiologists, for the purpose of propagation one pollen cell and one egg cell unite in Phanerogams into a single cell, which is capable by assimilation and formation of new cells, of becoming an independent organism ’’. However, this opinion was not generally accepted. Naudin (1863) repeated experiments of Kélreuter and Gartner, placing counted numbers of pollen grains on stigmas and concluding that a fully viable seed required more than one grain. Mendel himself repeated this experiment, using Mirabilis as had Naudin, and found, as he wrote in a letter to Nageli, that a single grain was sufficient. The foundations of our knowledge of chromosomes, mitosis and meiosis were laid between 1882 and 1885; Mendel died in 1884. Roux published a remarkable essay in 1883 in which he argued that the linear structure of the chromosomes and their precise longitudinal division were such striking and universal phenomena as to indicate selective value. He suggested that this lay in their effectiveness in assuring that each daughter cell received the same complement of chromosomal material. Further he regarded this as a strong argument for identifying the chromosomes as the bearers of the units of heredity. These units were here for the first time specified as being arranged in a linear series. We have no information as to whether Mendel himself considered his different pairs of factors as being other than independent from one another. In discussing the centenary of Mendel’s work, it has been proper to consider what Mendel did discover, how he did it and how he proved his theory. But my main purpose is to give some account of where genetics has now reached and the paths by which it has advanced. However, the advance has been so fast and so far that it is impossible to do justice to all of it. Particular attention will be paid to the nature of the hereditary factors and how they act, with a glance at possible future advances. The rediscovery in 1900 of Mendel’s work, independently by Correns, Tschermak and de Vries, was rapidly followed by evidence of its universal applicability to all organisms as a strong probability. About a quarter of a century or so later this evidence was extended to microorganisms, first to fungi and algae and later to bacteria and viruses. These extensions have resulted in remarkable advances in knowledge at an unprecedented rate. Early this century sex linkage, linkage and gene interaction were found as exceptions to the simple Mendelian rules. Polysomic inheritance, the effect of structural changes (such as interchanges, inversions and deletions) led to proofs that the chromosomes, visible microscopically, were the bearers of heredity. Probably the most fundamental of these discoveries was linkage. Many factors were found to be correlated in their segregation, rather than independent as Mendel’s Second Law states, with recombinants being in a minority. All experiments have agreed with the hypothesis that the factors belong to a number of distinct linkage groups and that those in a linkage group are arranged in a linear array. The hereditary material, which carries the factors or is composed of them, is filamentous. The name ‘ gene’ was introduced and used in the sense of Mendel’s factors in the germ cells. Genes were defined physiologically, namely that there is a heritable factor capable, under suitable circumstances, of being expressed as a recognisably distinct character. However, a gene responsible for a given character is detectable, in general, only if some other individual organism has AA 104 SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1966 a different, alternative gene responsible for a difference in the character. These two genes are allelic genes. Alleles form a set of genes all affecting one function in various ways. Whether two gene changes (mutations) are allelic or not is generally settled by the coupling and repulsion test, now often called the cis-trans test. Such changes in nomenclature, and there are many, as often reflect an ignorance of earlier knowledge as a radically changed point of view. Almost all mutational changes from the normal involve loss or partial loss of some characteristic. When combined with the normal, as a diploid in an animal or higher plant or in a yeast or as a heterocaryon in Neurospora or as a mixed population of bacteriophages growing in a bacterium, the mutant type is hidden by the normal. The normal is commonly dominant to the mutant which is recessive : it is as though the presence of a function in one covers up its absence in the other. Thus the problem of defining a gene as a unit of function is essentially that of deciding whether any two mutants are different changes of the same normal gene or changes of two different genes. If the same function were lost in both mutants, there would be no gain of function by their combination in a diploid or a heterocaryon. However, if different functions were lost in each mutant, the two would be able to complement each other’s loss, so restoring function. Thus with mutational changes in nonallelic genes, the coupling and repulsion phase heterozygotes both possess the normal function. However, with mutational changes in allelic genes, the repulsion phase heterozygote appears mutant whereas the coupling phase appears normal. In various microorganisms, it has been shown that, if a large number of mutants are tested in pairs, they are divisible into a number of groups, such that complementation always occurs between members of different groups. Within groups, either no complementation occurs between any pair or no complementation occurs between some of the individuals and all other members of the group, some pairs of which complement. In the latter case, the class of noncomplementing members holds the group together. The genes responsible for differences within a group are allelic while those responsible for differences between groups are nonallelic. The significance of complementation between alleles will be mentioned later. The differences between alleles, defined physiologically, are usually capable of genetic recombination. This has been demonstrated in Drosophila, maize, various fungi, bacteria and bacteriophages. Different pairs of alleles show different frequencies of recombination and these frequencies have been used to construct fine structure maps on the principle that the sites of mutation should be linearly arranged, with the frequency of recombination a function of the distance between them. The clearest and most extensive evidence is drawn from studies of the rlI genes of the bacteriophage T4. Here proof of the linearity is based on topological criteria rather than the statistics of recombination. Some r/Z mutants show no recombination with others, which themselves are able to recombine. The former can be regarded as more extensive abnormalities than the latter and may be deficiencies of various parts of the rIZ genes. If a series of them are analysed, some pairs showing recombination and others not, the results can be arranged in a matrix which can be represented as a linear map with the defects of the mutants extending over segments of various length. The overlaps define a unique order and none of the information requires the map to have more than one dimension. The total number of sites at which mutation has been shown to occur in rlI exceeds 300, a value which probably represents little more than a third of the possible sites. Thus, the heritable material is filamentous and is segmented into genes, distinguished by their functions. What are their functions? How do they D. G. CATCHESIDE 105: act? Of what are they made? What separates them? These are not independent matters, but certain functions were discovered first. Garrod, then Regius Professor of Medicine at Oxford University, suggested in 1909 that the function lost in a mutant was the ability to make a simple chemical transformation and that this was because of the loss of a specific enzyme. Hence, the proposition that each normal gene determined a specific enzyme. Garrod’s theory was based upon the study of inborn errors of metabolism in: man such as the failure in aleaptonuria to break down homogentisic acid and the failure in phenylketonuria to oxidise phenylpyruvie acid. Each of these metabolic defects is inherited as a simple recessive. This gene-enzyme theory was ignored for more than 30 years. In the meantime, Beadle and Tatum had made the same proposal in 1941 and were able to obtain a large series of biochemical mutants in the fungus Neurospora. Some of these mutants provided the first evidence of definitive changes in the properties of specific enzymes accompanying alterations in specific genes. Later it was shown that gene mutation was associated with changes in the composition of the protein of which enzymes are composed. Gradually, through the applica- tion of analytical procedures suitable for small amounts of complex materials, mainly due to advances in chromatographic separation of amino acids and peptides, it was shown that a given protein is made up of a particular number: of amino acids arranged in a precise sequence. It was shown also that the corresponding proteins, determined by heritable differences, may differ merely in the substitution of one amino acid for another at a definite place in the sequence. This was first shown for haemoglobin in man by Ingram in 1957, and has been extended to other systems, especially in bacteria and bacteriophages, by many workers. Among them, Yanofsky has shown, for the A protein of tryptophan synthetase in Escherichia coli, that the order of sites of allelic difference in the gene map parallels the order of positions of amino acid substitution in the protein specified by.the normal gene at the locus. Another demonstration of this parallelism, or congruence, due to Brenner and his associates, is dependent upon the fact that some kinds of mutation result in the formation merely of fragments of the normal protein. In bacteriophage 14, they were able to show a strict correlation between the position of the mutant. site in the gene map and the length of the peptide formed and so the position of discontinuity with reference to the normal polypeptide. Questions left to answer at this stage include how the gene determines. the protein, how the gene duplicates and what is the chemical and physical nature of the gene. It was a major achievement of the combined use of genetical and cytological methods to demonstrate that the chromosomes were the site of the genes. Further it was possible to demonstrate that there is a congruence in order of the genes in a linkage map and of the order of parts of a particular chromosome. Here it is reasonable to assume a collinearity in the sense that genes are parts of chromosomes, or of a constituent. The chromosomes were found to be composed of deoxyribonucleic acid (DNA), ribonucleic acid (RNA) and proteins, mostly basic ones. Amongst these materials, only the DNA was. in constant quantity from cell to cell of an organism, doubling before each mitosis. This was circumstantial evidence for DNA being the heritable material, but there was some reluctance to accept that it could show enough variety to be genetic, especially as it was thought to be merely a repeating sequence of the four nucleotides. At the same time most of the protein of the chromo- somes appeared equally monotonous, but some hope was centred in the small amounts of acidic protein present. Clear evidence for DNA being the genetic material of bacteria and of bacteriophages came first from the demonstrations that it was the material of transformation in Pneumococcus and virtually the only material injected by phages into host bacteria. In the meantime, DNA and RNA were subjected to thorough chemical study, so that by 1951 Todd and Brown had established 106 SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1966 in detail the structure of nucleosides and nucleotides and how they were linked together in nucleic acids to form linear molecules in which nucleotides are joined together by phosphate links to the 3 and 5 positions of their sugar residues. X-ray diffraction studies of DNA fibres enabled Wilkins to establish the constancy of pattern and approximate dimensions. Improvements in analysis of DNA from many sources resulted in the discovery of variations in the amounts of the four nucleotides, but with certain regularities, especially equivalence in the amounts of adenine and thymine and of guanine and cytosine, respectively, among the bases. To account for these data, Watson and Crick in 1953 proposed that DNA consists of two spiral polynucleotide chains, relationally coiled about one another and joined by hydrogen bonds between their bases. This structure is now generally accepted as correct in principle, subject to refinements in details of bond angles and atom spacing. Besides suggesting how the molecule might replicate and preserve the same genetic information in the daughters, it provided a mechanism of mutation and suggested how the information in the DNA might be related to that in the polypeptide of a protein. The double spiral structure predicts a semi-conservative mode of replication. Meselson and Stahl showed this to occur in the replication of DNA in Escherichia coli by following the distribution of heavy isotopes of nitrogen and carbon in the DNA of progeny developing in the presence of normal isotopes. Similar observations have been made with an alga and with mammalian cells. Labelling experiments to study the replication of chromosomes, in higher plants showed a similar semi-conservative process, but further work has disclosed complexities. Labelling experiments with the chromosomes of bacteria have also shown a semi-conservative process and, in addition, that the chromosomes are circular and are replicated sequentially starting from a particular site. How is DNA related to protein? These are two filamentous materials, one composed of four different members (the nucleotide pairs), the other of about 20 (the amino acids). It appeared possible that the relation was analogous to two different codes, one with four and the other with 20 symbols, for the same information. Obviously a one to one or a two to one relation gives insufficient scope. The least proportionality likely is a three nucleotide pair to one amino acid relation. It is unlikely that the code could be mixed, since it would require commas between the elements of the code as well as stops at the ends of the section of DNA specifying one protein. Also, it is unlikely that the code is overlapping, since then several adjacent amino acids would generally be affected by one mutation. The triplet code provides 64 symbols, some of which represent signals other than amino acids. Moreover, it has been found that some amino acids are represented by more than one triplet, a degeneracy of the code. The three to one relation was established for bacteriophage by ingenious experiments of Crick, Brenner and their associates. It was thought that certain mutants, induced by acridines, were due to the addition or subtraction of base pairs. Some reversions of these might have compensatory subtractions or additions near, but not at, the original mutant site. These suppressor mutations were separated and found themselves to be mutant. Whether base pairs were added or subtracted could be defined in terms of a standard. When three additive mutants, or three subtractive mutants, were put together the strain was approximately normal, rather than mutant, provided the sites of the mutants were fairly close together with specified parts of the gene. The significance of a third important constituent of living matter, RNA, became clear through a different sequence of events, particularly through the efforts of biochemists to determine the mechanism of protein synthesis. It is not necessary to go into details. In essence, it is now fairly well established that an RNA polymerase catalyses the synthesis of specific RNAs by trans- cription from segments of the DNA, so that the sequence in a given RNA is D. G. CATCHESIDE 107 the complement of the sequence in the strand of DNA copied. There are three classes of RNA molecules: (1) two species of ribosomal RNA (rRNA), one about 1,500 and the other 3,000 nucleotides long, that form parts of the structure of the ribosomes concerned in protein synthesis; (2) several dozen species of transfer RNA (tRNA), each about 100 nucleotides long, that provides adapters for the amino acids joined in protein synthesis; (3) hundreds of thousands of species of messenger RNA (mRNA), of diverse lengths, perhaps ranging up to tens of thousands of nucleotides, that provide the templates for orderly polymerisation of amino acids into specific polypeptides. Thus some genes determine only RNA (rRNA and tRNA) but most are expressed as protein. Each species of transfer RNA becomes coupled to a specific amino acid and the latter is led to its proper position in the amino acid order by a specific triplet in the tRNA complementary to a triplet in the messenger RNA. The discovery that polyribonucleotides stimulate the incorporation of amino acids in cell free systems able to synthesise proteins, led very quickly to solutions of which nucleotide triplets correspond to each amino acid. This was achieved by using synthetic polynucleotides composed either of single nucleotides or of random mixtures of two or more in various proportions. Of course, the information is incomplete and it is not yet known exactly how many of the 64 triplets do code for amino acids. Some triplets do not code for any amino acid and these have been referred to as nonsense triplets. In fact this is an unsuitable name, since at least some of them serve a special function in information transfer. In bacteria and bacteriophages, it has been shown that some mutations result in the termination of synthesis of a polypeptide so that only a fragment of the normal one is formed. Termination occurs because the mutation has resulted in triplet which signals that this very event should occur. The evidence is strictly genetical, dependent upon the demonstration that a distinct suppressor mutation may allow an amino acid to be inserted despite that the signal normally leads to the cessation of amino acid polymerisation. Thus we have a strictly mechanistic theory of gene structure, function and replication. The gene is a sequence of nucleotide pairs, presumably a multiple of three in number. The gene replicates by enzymic processes and is transcribed into RNA by another enzymic process. Most species of RNA provide templates, when wound on a number of ribosomes, for ordering specific transfer RNAs, to which are coupled acylated amino acids, which can then be bound together in an order, specified by the messenger RNA. The polypeptides so formed fold to form proteins, generally enzymes. Many enzymes are composed of several similar protein units and in these cases some defective allelic mutants may complement in a hybrid enzyme by allosteric correction of folding errors in the mutant proteins. The theory is further that there is a gene or a complex of genes for each and every function in an organism. This is substantiated by the discovery of genes for each enzyme of intermediary metabolism, of genes for some parts of the machinery of protein synthesis, of genes for a wide variety of morpho- logical, physiological and behavioural characters, of genes for behaviour of constituents of cells, especially the chromosomes and mitochondria, and of genes controlling the action of other genes. Genetics is a powerful method of biological analysis, discovering principles and mechanisms by analysis of numerical data of the distribution of heritable differences amongst progeny. In essence its principles extend from bacterio- phage to man; it unifies biology. It may be expected soon to make decisive contributions to some of the outstanding fundamental problems in biology, such as the mechanisms of genetic recombination, differentiation and behaviour. Mendel did indeed give us a legacy, one I am sure Sir William Macleay would appreciate were he alive today to see how genetics has made such good intellectual use of the bacteria, whose general study he fostered. I am grateful for the opportunity afforded of paying respectful tribute to both these benefactors. 108 SIR WILLIAM MACLEAY MEMORIAL LECTURE, 1966 Bibliography GENERAL REFERENCES The following deal with broad aspects of the subject and contain references to nineteenth century work— Intis, H., 1924.—‘‘ Gregor Johann Mendel, Leben, Werk und Wirkung”’. (Springer, Berlin). English translation, E. and C. Paul, 1932, “ Life of Mendel”. (W. W. Norton and Company, New York). Roserts, H. F., 1929.—** Plant Hybridization before Mendel”. (Princeton University Press, Princeton, N. J.). Sacus, J., 1875.—‘* History of Botany’’. Translated by H. E. F. Garnsey, 1890. (Oxford University Press, Oxford). SturtTEvANT, A. H., 1965.—“* A History of Genetics”. (Harper and Row, New York). Witson, HE. B., 1896.—* The Cell in Development and Heredity ”. (Macmillan Co., New York). Second edition, 1900; Third edition, 1925. ZIRKLE, C., 1935.—“ The Beginnings of Plant Hybridization’”’. (University of Pennsylvania Press, Philadelphia). SPECIAL REFERENCES BEADLE, G. W., and Tatum, HE. L., 1941.—Genetic control of biochemical reactions in Neurospora- Proc. nat. Acad. Sci., 27: 499-506. Benzer, S., 1961.—On the topography of the genetic fine structure. Proc. nat. Acad. Sci., 47: 403-415. BRENNER, S., 1966.—Collinearity and the genetic code. Proc. roy. Soc., B164: 170-180. Brown, D. M., and Lytueor, B., 1950.—Deoxyribonucleosides and related compounds. Part II. A proof of the furanose structure of the natural 2-deoxyribonucleosides. J. chem. Soc., 1990-1991. Crick, F. H. C., Barnett, LESLIE, BRENNER, S., and Warts-Topin, R. J., 1961.—General nature of the genetic code for proteins. Nature, 192: 1227-1232. CoRRENS, C., 1900.—G. Mendel’s Regel tiber das Verhalten der Nachkommenschaft der Rassen- bastarde. Ber. deutsch. bot. Gesells., 18: 158-168. English translation 1950, in The birth of genetics. Suppl. Genetics, 35: 33-41. , 1905.—Gregor Mendel’s Briefe on Carl Nageli, 1866-1873. Abh. math.-phys. Kl. Kén. Sdchs. Gesellsch. Wiss., 29: 189-265. English translation, 1950, in The birth of genetics. Suppl. Genetics, 35: 1-29. DeE Vrizs, H., 1900.—Sur la loi de disjonction des hybrides. C. R. Acad. Sci. (Paris), 130: 845-847. English translation 1950, in The birth of genetics. Suppl. Genetics, 30-32. FisHer, R. A., 1936.—Has Mendel’s work been rediscovered? Ann. Sci., 1: 115-137. Garrop, A. E., 1908.—Inborn errors of metabolism. Lancet, 2: 1—7, 73-79, 142-148, 214-220. Published separately as a book, 1909. (Oxford University Press, London). Ineram, V. M., 1963.—** The Hemoglobins in Genetics and Evolution’. (Columbia University Press, New York). LytHeor, B., and Topp, A. R., 1947.—Structure and synthesis of nucleotides. Symp. Soc. exp. Biol., 1: 15-37. MENDEL, G., 1866.—Versuche tiber Pflanzenhybriden. Verh. naturforsch. Ver. Brunn, 4: 3-47. English translations, 1901. J. roy. hort. Soc., 26: Bateson, W., 1909, 1913. ‘* Mendel’s principles of Heredity ”’. Mesetson, M., and Stan, F. W., 1958.—The replication of DNA. Cold Spring Harbor Symp. quant. Biol., 23: 9-12. TscHERMAK, H., 1900.—Uber ktnstliche Kreuzung bei Pisum sativum. Ber. deutsch. bot. Gesells., 18: 232-239. English translation 1950, in The birth of genetics. Suppl. Genetics, 35: 42-47. Watson, J. D., and Crick, F. H. C., 1953.—The structure of DNA. Cold Spring Harbor Symp. quant. Biol., 18: 123-131. YANOorFSKyY, C., Cartton, B. C., Guest, J. R., Henrsxki, D. R., and Hennine, U., 1964.—On the colinearity of gene structure and protein structure. Proc. nat. Acad. Sci. U.S., 51: 266-272. THE RHAPHIDOPHORIDAE (ORTHOPTERA) OF AUSTRALIA 4. A NEw GENUS FROM SouTH AUSTRALIA AoLA M. RICHARDS Department of Zoology, University of New South Wales, Sydney {Read 29th June, 1966] Synopsis A new genus Novotettix n.g. is erected, and the new species Novotettix naracoortensis n.sp. is described from limestone caves in south-eastern South Australia. INTRODUCTION A new monotypic genus Novotettix n.g., belonging to the family Rhaphidophoridae, is recorded here from Naracoorte in south-eastern South Australia. The new species, Novotettix naracoortensis n.sp., is placed in this genus. The caves at Naracoorte are formed in Oligocene Gambier marine limestone, and occur at an altitude of about 200 feet above sea level. Twenty-eight caves are known, covering an area of 15 miles (Sexton, 1965). N. naracoortensis has been collected from Alexandra Cave and Victoria Cave on the South Australian Tourist Bureau Reserve. Specimens have also been taken in Haystall Cave, Corner Fence Cave, and Smoke Cave at Joanna, a district 9-10 miles south-south-east of the town of Naracoorte, and about four miles from the Tourist Reserve. ‘ In both Alexandra and Victoria Caves temperatures range from 62°-64°F and relative humidity from 89-94°,. Similar temperatures have been recorded from caves containing Rhaphidophorids in western Victoria and on the Nullarbor Plain. These temperatures are considerably higher than those recorded from caves in south-eastern Australia, where the range is from 49°-58°F, and the average temperature 54°-56°F (Richards, 1966). At Waitomo Caves in New Zealand, the highest temperature at which Rhaphidophorids could carry out their normal activities was found to be 60°F (Richards, 1965). Average cave temperatures there ranged from 53°-58°F. The southern Australian caves are the warmest habitats so far recorded in Australasia which support colonies of Rhaphidophoridae. The higher temperatures may be attributed to the low altitudes at which the caves occur. Alexandra Cave contains a colony of over 1500 N. naracoortensis, in contrast to the rather sparsely populated caves in south-eastern Australia. About two-thirds of the insects occur on the walls 100 feet inside the entrance, and the remainder extend another 70 feet or more into the cave. The Rhaphido- phorids share the cave with a fauna of carabids, pterostichids, isopods, two species of blattids, and two species of spiders. The very large number of Rhaphidophorids present may be accounted for by the absence of predators such as bats and rats. The Bat Cave, a short distance from Alexandra Cave, has a large colony of the bent-winged bat, Miniopterus schreibersii (Kuhl), but no NV. naracoortensis have been observed in this cave. In Tasmanian caves, where no bats occur, large colonies of Rhaphidophoridae are also present. This dissociation of cave crickets and bats is quite the exception to the rule in caves throughout Australia. PROCEEDINGS OF THE LINNEAN SocreTy or New SoutH Watss, Vol. 91, Part 2 110 RHAPHIDOPHORIDAE (ORTHOPTERA) OF AUSTRALIA N. naracoortensis is one of the larger of the Australian Rhaphidophoridae. Sexual dimorphism is strongly developed, males being larger than females. An adult male may reach a length of up to 18 cm. from the tip of its antennae to its hind tarsi. The life history follows a similar pattern to that observed in other Rhaphidophorids, both in Australia and New Zealand (Richards, 1961, 1966). In late December and January adults appear. Mating and oviposition continue throughout the autumn. The locomotor activity rhythm of N. naracoortensis has recently been discussed, and the similarity of its behaviour to that of New Zealand species noted (Richards, 1965). This new genus Novotettix does not show any close affinities with other Australian Rhaphidophorid genera so far studied. Genus NOVOTETTIX, n.g. Body sparsely clothed with short setae. Legs long and slender. Antennae very long and tapering, almost touching at their bases; scape about three times as large as pedicel, which is narrower than scape, but broader than other segments ; from segment four onwards segments subequal in length, but steadily decreasing in size; all segments thickly clothed with short setae. A single median ocellus present. Fastigium rising abruptly, grooved medianly and longitudinally. Maxillary palps with third and fourth segments subequal in length. Fore coxa unarmed. All femora sulcate ventrally. Apical spines on femora, tibiae, proximal segments one and two of hind tarsi constant in number. Fore femur bears two apical spines beneath, one prolateral and one retrolateral ; fore tibia bears four apical spines, one above and one beneath both prolaterally and retrolaterally ; fore tarsus unarmed. Mid femur bears two apical spines beneath, one prolateral and the other retrolateral ; mid tibia bears four apical spines, one above and one beneath both prolaterally and retrolaterally ; mid tarsus unarmed. Hind femur bears one prolateral apical spine beneath ; hind tibia bears a pair of long apical spurs above, a pair of subapical spines above, a pair of short apical spurs beneath and a pair of subapical spines beneath, one from each pair being prolateral and the other retrolateral; two proximal segments of hind tarsus each bear two apical spines above, one prolateral and one retrolateral; other two segments unarmed. Subgenital plate of female trilobed, each lobe rounded apically. Subgenital plate of male triangulate. Type species for the genus: Novotettix naracoortensis n.sp. VOVOTETTIX NARACOORTENSIS, 0.Sp. (Text-fig. 1. Figures 1-6) Colour.—Basic colour ochreous with pronotum, mesonotum, metanotum and abdominal terga irregularly mottled with light brown ; femora and tibiae banded or mottled with light brown and ochreous; all tarsi ochreous; antennae light brown ; oOvipositor light reddish brown. Body.—Length up to 16 mm. in male, and 17 mm. in female. Surface of body sparsely clothed with setae. Antennae broken. Fastigium longer than high. Ovipositor 0-9 length of body; ventral valves armed distally 0-3 of total length to apex with seven well developed teeth (Fig. 1). Antennae.—As in generic description. Third segment in male on dorsal aspect 2-6 as long as pedicel, and on ventral aspect 2-1 as long; in female on dorsal aspect 1:6 as long as pedicel, and on ventral aspect 1-3 as long. Sexual dimorphism present, male possessing longer, stouter antennae than female. No spines present on flagellum of either male or female. AOLA M. RICHARDS mint Legs.—Fore and mid legs subequal in length, with hind leg 1-7 length of fore and mid legs. Sexual dimorphism is shown by fore, mid and hind legs of female being 0-8 as long as male. Hind femora, all tibiae and proximal two segments of hind tarsi armed with variable numbers of linear spines Text-fig. 1—DNovotettix naracoortensis n.sp. 1, Distal portion of ovipositor showing teeth on ventral valve; 2, Female genitalia, dorsal view; 3, Female genitalia, ventral view; 4, Male genitalia, dorsal view; 5, Male genitalia, ventral view; 6, Male genitalia, ventral view, subgenital plate removed to expose structures beneath. 112 RHAPHIDOPHORIDAE (ORTHOPTERA) OF AUSTRALIA (Table 1). No spines on fore or mid femora and tarsi. Spines on hind femur of female fewer in number than those on hind femora of male. Apical spines constant in number as in generic description. Length of proximal segment of hind tarsus subequal with other three segments together. Ratio of length of legs to length of body: Fore leg, male 2:6:1; female 2:1. Mid leg, male 2-6:1; female 2:1. Hind leg, male 4:6:1; female 3-3: 1. TABLE | Variability in Number of Linear Spines on the Legs of 25 Specimens of Novotettix naracoortensis n.sp. Arith. No. of Std. Range Mean Specimens Dev. (or Distribution) L. R. 1h IR. Th, R. L. R. Fore Femur Pro. 0 0 25 25 0 0 0 0 Inf. Retro. 0 0 25 25 0 0 0 0 Fore Tibia Pro. 3:2 3:2 25 25 0:6 0-6 2-4 2-4 Inf. Retro. 4-0 4-0 25 25 - - 3(2), 4(23) 3(2), 4(23) Fore Tarsus Pro. 0 0 25 74)5) 0 0 0 0 Retro. 0 O- 25 25 0 0 0 0 Mid Femur Pro. 0 0 25 25 0 0 0 0 Inf. Retro. 0 0 25 25 0 0 0 0 Mid Tibia Pro. ) 0 25 25 0 0 0 0 Sup. Retro. 0 0 25 25 0 0 0 0 Mid Tibia Pro. 3:7 3:7 25 25 0:5 0-5 3(7), 4(18) 2-4 Inf. Retro. 4-0 3°8 25 25 0:3 0:5 3-5 3-5 Mid Tarsus Pro. 0 0 25 25 0 0 0 0 Retro. 0 0 25 25 0 0 0 0 Hind Femur Pro. 13-0 14:1 14 14 2-4 3°1 9-17 8—21 Inf. ¢ Retro. 16-1 17-9 14 14 4-0 3:6 9-25 13-28 Hind Femur Pro. 7-7 8-1 11 10 2-7 2-0 5-13 4—12 Inf. 2 Retro. oR 7:8 11 10 2-4 2-3 3-11 4-1] Hind Tibia Pro. 63-6 62-6 25 24. 4-1 6-6 57-72 48-75 Sup. Retro. 68-9 67-7 25 24 8-1 6:6 55-81 59-83 Hind Tarsus Pro. 8:8 8-4 25 24 1-2 1:3 6-11 6-11 1 Sup. Retro. 8:8 8-5 25 24 1:9 1:6 6-15 6-11 Hind Tarsus Pro. 3-1 3:0 25 24 0:7 0:7 2-5 2-4. 2 Sup. Retro. 2°8 3-0 25 24 0-6 0:8 2-4 2-5 Arith. Mean, Arithmetic Mean; Inf., Inferior; L., Left leg; Mid., Middle; Pro., Prolateral ; R., Right leg; Retro., Retrolateral; Std. Dev., Standard Deviation ; Sup., Superior. (Figures in parentheses represent number of specimens) Genitalia. FEMALE: Suranal plate, Fig. 2 (SAP), concave laterally, distal margin slightly emarginate ; whole plate thickly clothed with setae. Subgenital plate, Fig. 3 (SGP), distal margin trilobed, each lobe rounded apically, two lateral lobes 0-25 longer than median lobe; whole plate sparsely clothed with setae. MALE: Suranal plate, Fig. 4 (SPL), convex laterally, distal margin emarginate and clothed with setae; rest of plate thickly clothed with setae on dorsal and ventral surfaces ; on ventral surface disto-lateral regions slightly raised into two lobes. Subgenital plate, Fig. 5 (H), triangulate, 2-5 wider than long, sparsely clothed with setae. Two styli, Fig. 5 (S), short, broad, conical, thickly clothed with setae, length of styli being 0-7 length of sternite IX (S IX). Parameres, Figs 5, 6 (P), elongate, tapering to a point, retrolateral AOLA M. RICHARDS 113 margin concave, prolateral margin convex, whole plate 2-7 longer than wide, distal portion of paramere thickly clothed with long and short setae. Pseudo- sternite, Fig. 6 (PD), 1-5 wider than long, convex laterally, tapering distally to a truncated apex. Penis, Fig. 6 (PN), two-lobed, each lobe subequal in width to length. Paraprocts, Figs 5, 6 (PP), crescent-shaped, twice as long aS wide, sparsely clothed with setae. Locality.—In limestone caves, Naracoorte, South Australia. Alexandra Cave (S4) (type locality), coll. A. M. Richards, 1964; G. F. Gross, 1958; E. Hamilton-Smith, 1956, 1958, 1961, 1962; MHaystall Cave (S34), coll. P. F. Aitken, 1962 ; Smoke Cave (S871), coll. P. F. Aitken, 1962 ; Corner Fence Cave (S35), coll. P. F. Aitken, 1962; Victoria Cave (S2), coll. W. Hill, 1957; Cave two miles east of Naracoorte, coll. unknown, 1957. Types.—Holotype male, allotype female, and two paratypes (male and female) in National Insect Collection, C.S.I.R.O., Canberra. Four paratypes (two males and two females) in South Australian Museum Collection, Adelaide. Two paratypes (male and female) in Australian Museum Collection, Sydney. Acknowledgements I am indebted to Dr. P. Crowcroft, Director of the South Australian Museum, Adelaide, for the loan of material. I should also like to thank Mr. E. Hamilton-Smith, Sub-Aqua Speleological Society, Melbourne; and Mr. P. F. Aitken, South Australian Museum, for collecting and sending me specimens. I am grateful to the South Australian Government Tourist Bureau for permission to collect and study insects in the caves on their Reserve at Naracoorte. References Ricuarps, Aola M.. 1961.—The life history of some species of Rhaphidophoridae (Orthoptera). Trans. roy. Soc. N.Z. Zoology, 1 (9): 121-137. , 1965.—The effect of weather on Rhaphidophoridae (Orthoptera) in New Zealand and Australia. Ann. de Spéléologie, XX- (3): 391-400. , 1966.—The Rhaphidophoridae (Orthoptera) of Australia. Part 3. A new genus from south-eastern Australia. Pacific Insects, 8 (3): 613-624. Sexton, R. T., 1965.—Caves of the coastal areas of South Australia. Helictite, 3 (3): 45-59. Index to Text-figure BC, basal segment of cereus; C, cereus; DV, dorsal valve; EP, endoparamere ; H, subgenital plate, male; IA, intersegmental apodeme; MT IX, membrane of tergite IX; P, paramere (ectoparamere) ; P VIII, P IX, pleurite VIII, [IX ; PD, pseudosternite ; PM, perianal membrane ; PN, penis; S, stylus; S VII, S VIII, S IX, sternite VII, VIII, IX ; SAP, suranal plate, female ; SGP, subgenital plate, female; SPL, suranal plate, male; T VII, T VIII, T IX, T X, tergite VII, VIII, IX, X; 1 VF, first valvifer; 2 VF, second valvifer; VV, ventral valve. NOTES ON THREE RECENTLY PROPOSED AUSTRALIAN TERTIARY ECHINOID GENERA G. M. PHILIP Geology Department, University of New England [Read 29th June, 1966] Synopsis The Eocene brissid genus Gillechinus Fell, 1964 (type species: Giullechinus cudmorei Fell, 1964=Hupatagus coranguintum Tate MS.) is considered to be a synonym of Brissopatagus Cotteau. 1886. The Miocene temnopleurid genus Irenechinus Fell, 1964 (type species: Irenechinus hentyr Fell, 1964=Coptechinus pulchellus Bittner, 1892) is considered to be a synonym of Ortholophus Dunean, 1887. Brochopleurus australiae Fell, 1949 (=Psammechinus woodsi Laube, 1869) should also be referred to Ortholophus. The Eocene fibulariid Lenicyamidia Brunnschweiler, 1962 (type species: Lenicyamidia compta Brunnschweiler, 1962) is considered to possess a typical clypeasteroid monobasal apical system. In this note three recently proposed Australian Tertiary echinoid genera are discussed. Although the Tertiary echinoid faunas of Australia are at present being revised, it will be several years before relevant sections dealing with all these genera appear. In the interim it is desirable that some comments be given before these genera become established in the literature. Genus GILLECHINUS Fell, 1964 This genus was proposed for a species of spatangoid from the ‘‘ Lower beds, Aldinga, South Australia”? which was named Gillechinus cudmoret by Fell. The species is in fact Ralph Tate’s MS. form Hupatagus coranguinium and has been cited as such in literature on the Australian Tertiary (e.g. Pritchard, 1891, p. 183). In the collections available to me (including Tate’s original syntypes from the Tate Collection, University of Adelaide) it appears that the species is confined to the Upper Eocene Tortachilla Limestone of the St. Vincent Basin, South Australia. According to Fell, Gillechinus is similar to Hupatagus but differs ‘‘in the arrangement of the primary tubercles” and the ‘indistinct closure of the petals”. He goes on to observe that Gillechinus is unknown outside Australia. However, in the comparison of the genus with previously described forms, no mention is made of Brissopatagus Cotteau, 1863 (type species: Brissopatagus caumonti Cotteau). This brissid is world-wide in its distribution (Europe, Egypt, Madagascar, India, N. America) and is confined to Eocene strata. It differs from Hupatagus solely in the tuberculated, somewhat depressed areas anterior to each petaloid ambulacrum of the adapical surface. In some species these depressions may be present only in front of the anterior petals, whereas in others the depressions are not well defined. Indeed, Cooke (1942, 1959) has employed Brissopatagus as a subgenus of Hupatagus, for the differences from Hupatagus hardly warrant its generic separation. Gillechinus cudmorei Fell is a typical species of Brissopatagus. Among previously described forms embraced by this genus it resembles most closely Euspatangus (Brissospatangus) beyrichi Dames (1877, p. 82-83, Pl. 11, Fig. 2) PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vol. 91, Part 2 G. M. PHILIP 115 from the Eocene of Verona. It differs from this form in its smaller size, and wider, more rounded outline. Gillechinus Fell, 1964, therefore, is here considered to be a synonym of Brissopatagus Cotteau, 1886. Genus IRENECHINUS Fell, 1964 Fell (1964) has proposed the name Irenechinus hentyi gen. et sp. nov. for a specimen of a temnopleurid from the Middle Miocene Bochara Limestone, Hamilton district, Western Victoria. The elucidation of this genus requires discussion of some other Tertiary temnopleurids, in particular, Brochopleurus australiae Fell and other species which should be embraced by the genus Ortholophus Duncan. Fell (1949) described the temnopleurid Brochopleurus australiae sp. nov. from early Miocene strata exposed along the Murray River Cliffs, South Australia. Examination of many hundreds of specimens from these strata indicates that the holotype (National Museum Victoria P4687) of this species is a juvenile specimen of the common temnopleurid ‘‘ Psammechinus ”’ woodsi Laube, 1869. The species exhibits a progressive loss of sculpture during growth so that in large specimens the ridges tend to meet and the sculpture is correspondingly obliterated. A similar loss of sculpture during growth has been reported in living temnopleurids such as Desmechinus and Pseudechinus. It follows, therefore, that Brochopleurus australiae Fell, 1949 is a junior subjective synonym of ‘‘ Psammechinus”’ woodsi Laube, 1869. A second, poorly preserved specimen which Fell included in his species (National Museum Victoria P4688), is here interpreted as the male form of Paradoxechinus novus Laube. Such small specimens differ from ‘“ P.”’ woodsi principally in the character of the coarser sculpture. Fell’s generic assignment of the species to the Indian Miocene genus Brochopleurus also cannot be accepted. In 1887 Duncan proposed the genus Ortholophus for his species Temnechinus lineatus Duncan 1877 (non Bittner, 1892) from Upper Miocene strata at Beaumaris, Victoria. In the past there has been much uncertainty as to the application of this genus (e.g. Mortensen, 1943, pp. 350-1) due to discrepancies in Dunean’s descriptions. He gave his species as ‘‘ sculptured by a network of ridge-like costae” but his figure of O. lineatus shows no trace of sculpture. Examination of the type specimen (British Museum (Nat. Hist.) G.S.L. 14078), together with topotype material, shows that Duncan’s figure correctly portrays the ornament of mature specimens. In fact Ortholophus is a well founded genus and should embrace a group of Australian (and perhaps New Zealand) Tertiary temnopleurid species. The genus thus employed includes moderate sized temnopleurids with crenulate tubercles, uniserial ambulacra and obscure gill slits. The apical system is regularly dicyclic. Juveniles are strongly sculptured, the sculptural ridges being lost with growth to varying degrees in different species. The girdle consists of rather low auricles united above the ambulacra. Psammechinus woodst Laube is a typical representative of Ortholophus. Ortholophus differs from Brochopleurus principally in possessing crenulate tubercles, and hence Brochopleurus is not considered to be a junior synonym of Ortholophus. Fell (1949), however, described and illustrated his Brochopleurus australiae (a small specimen of Ortholophus woodsi) as possessing smooth tubercles. The holotype is not well preserved in this respect but distinct crenulation of the tubercles is readily discernible. In this respect it may also be noted that Bittner, Duncan and Laube have all erroneously described (and illustrated) the tubercles as smooth in various species of Ortholophus. To return to the genus Jrenechinus Fell, this is distinguished by Fell from Brochopleurus by the crenulation of the tubercles. It is in fact a typical species of Ortholophus in which Fell has recognized the crenulation of the tubercles. The species J. hentyi is also identical with that which occurs in the Middle 116 THREE RECENTLY PROPOSED AUSTRALIAN TERTIARY ECHINOID GENERA Miocene horizons along the Murray River Cliffs (the Morgan Limestone) and which was named by Bittner in 1892 (p. 342-4, Pl. 1, Fig. 5) as Coptechinus pulchellus. It is concluded that : Brochopleurus australiae Fell=Ortholophus woodsi (Laube) Irenechinus hentyi Fell=Ortholophus pulchellus (Bittner) The genus Brochopleurus is accordingly unknown from the Australian Tertiary. Illustrations of these and related temnopleurids will be given elsewhere (Philip, 1966). Genus LENICYAMIDIA Brunnschweiler, 1962 This clypeasteroid genus, based on L. compta Brunnschweiler, 1962, from the Eocene Merlinleigh Sandstone, Western Australia, was proposed for a coarsely tuberculated fibulariid with an elongate periproct. Particular interest attaches to the apical system of the type species, for it is described as consisting of ‘‘ Four large genital, one central madreporic, and five small ocular plates .. .” (Brunnschweiler, 1962, p. 167). Such an arrangement of plates is shown in Text-fig. 2A accompanying the original description. Fig. 1.—Lenicyamidia compta Brunnschweiler. Adapical portion of petals and apical system showing the fibrils of the stereome of the madreporite. Paratype, Bureau of Mineral Resources CPC 4812, x20 approx. All known representatives of the order Clypeasteroida possess a monobasal apical system, i.e., a fused genital disc (the madreporite), with five oculars which usually can be distinguished from the madreporite. The nature of the madreporite is known in Hchinarachnius where genitals 1 and 3 are lost at metamorphosis and genital 5 does not develop (Gordon, 1929). Genitals 2 and 4 alone fuse to give the madreporite. The apical system of Lenicyamidia compta would therefore seem to represent a remarkable departure from that of other clypeasteroids. Not only is the apical system of Lenicyamidia, as portrayed, unique among clypeasteroids, but with its ‘‘ central madreporiec plate ’’ (which would therefore be genital 2) it is without parallel in the class Echinoidea. Examination of type material of LZ. compta, however, indicates that the species possesses a monobasal system. All material of the species is silicified, and it was found impossible to develop sutures in the adapical region of any G. M. PHILIP 117 specimen. However, the silification has not obliterated the original fibrils of the stereome. These radiate from the central portion of the apical system to outside the genital pores without marked discontinuities (text-fig. 1). Where alignment of the fibrils of plates of an echinoid test is discernible, the fibrils are generally normal to sutures. It is therefore concluded that Lenicyamidia compta lacks discrete genital plates in the apical system which, in its nature, corresponds closely with that of other clypeasteroid echinoids. A comparison of Lenicyamidia with the closely related genera Fibularia, Fibulariella and Cyamidia will be given elsewhere. Acknowledgements I am obliged to Dr. N. H. Fisher, Bureau of Mineral Resources, Geology and Geophysics, Canberra, Mr. E. D. Gill, National Museum of Victoria, and Professor M. F. Glaessner, University of Adelaide, for the loan of specimens. Current work on Australian echinoderm faunas is generously supported by a University of New England Research Grant. References Birrner, A., 1892.—Uber Echiniden des Tertiars von Australien. Sitz. k. Adad. Wissen. Wien, (Math. Natur. Cl.), 101: 331-378. BRUNNSCHWEILER, R. O., 1962.—On echinoids in the Tertiary of Western Australia with a description of two new Eocene Fibulariidae. J. geol. Soc. Aust., 8: 159-169. Cooks, C. W., 1942.—Cenozoic irregular echinoids of eastern United States. J. Paleont., 16: 1-62. , 1959.—Cenozoic echinoids of eastern United States. U.S. geol. Surv. prof. Pap., 321: 1-106. Dames, W., 1877.—Die Hchiniden der vincentinischen und beronesischen Tertiaerablegerungen. Paldontograph., 25: 1-100. Duncan, P. M., 1877.—On the Echinodermata of the Australian (Tertiary) deposits. Quart. J. geol. Soc. Lond., 33: 42-71. , 1888.—A revision of the Echinoidea from the Australian Tertiaries. Jbid., 43 : 411-430. FEL, H. B., 1949.—An echinoid from the Tertiary (Janjukian) of South Australia, Brochopleurus australiae sp. nov. Mem. nat. Mus. Vict., 16: 17-19, fig. 1. , 1953.—The origins and migrations of the Australasian echinoderm faunas since the Mesozoic. Yrans. roy. Soc. N.Z., 81: 245-255. , 1964.—New genera of Tertiary echinoids from Victoria, Australia. Mem. nat. Mus. Vict., 26: 211-215. Gorpon, Isabella, 1929.—Skeletal development in Abracia, Echinarachnius and Leptasterias. Phil. Trans. roy. Soc. Lond., Ser. B., 215: 255-313. LauBs, G. C., 1869.—Uber einige fossile Echiniden von den Murray Cliffs in Sud-Australien. Sitz. k. Akad. Wissen. Wien. (Math. Natur. Cl.), 59: 183-198. Mortensen, Th., 1943.—*‘ A monograph of the Echinoidea”’ III (2), Camarodonta I. 533 pp. (C. A. Reitzel, Copenhagen). Purip, G. M., 1966.—The Tertiary echincids of south-eastern Australia. Part IV—Camarodonta (2). Proc. roy. Soc. Vict. (in preparation). PRITCHARD, G. B., 1891.—Catalogue of Australian Older Tertiary Mollusca and Pliocene species in South Australian School of Mines Museum. Ann. Rept. S.A. School of Mines and Industries, 176-206. CHROMOSOME NUMBERS IN LAMPROTHAMNIUM! A. T. HOTCHKISS Department of Biology, University of Louisville Louisville, Kentucky, U.S.A. 40208 [Read 29th June, 1966] Synopsis A comparison is made of the chromosome numbers of four species of Lamprothamnium, L. papulosum, L. succinctum, L. macropogon and Lamprothamnium sp., all polyploid, with the range of polyploid numbers in the related genus Chara. The possible use of chromosome number in the clarification of relationships and geographical distribution in Lamprothamnium is discussed. Lamprothamnium J. Groves (1916) is a small genus of the family Characeae distributed in the Hastern Hemisphere from Europe through Africa and southern Asia to the Australasian region. It occurs in brackish water, usually quite close to the sea. Despite a widespread distribution and abundant records, the genus remains obscure in its relationships, and the limits of some species are uncertain (Ophel, 1947; Wood and Imahori, 1965). As stated by Wood (Wood and Imahori, 1965) : The status of genus Lamprothamnium is not clear. The species form a natural group and as such are accepted as a genus. On the other hand, Lamp. succinctum exhibits certain Chara-like features, especially the occurrence of oogonia above the antheridia, and the bract-cells and stipu- lodes are not huge as is so common in many forms of Lamprothamnium. It seems that this series of species is more similar to Chara subgen. Charopsis than subgen. Charopsis is to subgenus Chara. Lamprothamnium could well be construed to be a third subgenus of genus Chara, and be considered as derived recently from subgen. Charopsis. Broadly based systematic studies are required for an evaluation of the position of Lamprothamnium in the Characeae. In the present. paper, reports of cytological investigations (Hotchkiss, 1963, 1965) are continued with collections of Lamprothamnium from Europe and Australia. Lamprothamnium papulosum (Wallr.) J. Groves was collected in Denmark on September 5, 1965 and kindly sent by T. Christensen. Specimens of Lamprothamniwm from the extensive TABLE | Chromosome Numbers Reported in the Genus Lamprothamnium Species with conjoined gametangia Lamprothamnium papulosum (Wallr.) J. Gr. ca. 50 Europe (Lindenbein, 1927) Lamprethamnium papulosum (Wallr.) J. Gr. 56 Denmark (present report) Lamprothamnium sp. 28 Western Australia (Hotchkiss, 1963, as L. macropogon) Species with disjunct gametangia Lamprothamnium succinctum (A. Br. in Asch,) Wood 42 New Caledonia (Hotchkiss, 1965) Lamprothamnium macropogon (A. Br.) Ophel 56 South Australia (present report) cytological collections of charophyte materials gathered by R. D. Wood in the Australasian region and described in preliminary reports by Wood (1962b) and Hotchkiss (1965), are as follows: WESTERN AUSTRALIA: Lamprothamnium sp., Yadgers Brook, 12 miles S of Moora, forming large clump at rock rapids. This study was supported in part by the National Science Foundation, U.S.A. Contribution No. 89 (New Series) of the Department of Biology, University of Louisville. PROCEEDINGS OF THE LINNEAN SocitETY oF NEw SoutH WaAtEs, Vol. 91, Part 2 A. T. HOTCHKISS 119 R. D. Wood 60-10-1-5, October 1, 1960; SourH AUSTRALIA: Lamprothamniwm macropogon (A. Br.) Ophel, in one ft. of water, on pipe clay, forming dense carpet under Ruppia (2), lagoon near shack, ca. one mi. E of Chinamans Well, 13 mi. SE of Salt Cr., Coorong region. R. D. Wood and C. von der Borsch 60-9-22-1, September 22, 1960. Chromosome number was determined in mitoses in antheridial filaments stained with orcein after fixation in acetic alcohol. Figs 1-4. Chromosome complements in Lamprothamnium present the appearance of numerous small to medium size chromosomes with about one larger chromosome in every 14 (Figs 1-4). The two new chromosome number determinations in the present report, together with those from the literature, are summarized in Table 1, an enumeration which shows the strongly polyploid nature of the taxa thus far investigated in this genus. DISCUSSION The base chromosome number in the Tribe Chareae of the Characeae appears to be 14. The new determinations reported here are in accord with this pattern and extend the known chromosome numbers in Lamprothamnium to the n=56 level. The euploid series of 14, 28, 42 and 56 chromosomes, well known in the large genus Chara, is repeated in the 28, 42, and 56 series found thus far in species of the much smaller genus Lamprothamnium. Tt is interesting that the estimate of n=ca. 50 for L. papulosum, in an early report by Lindenbein (1927) under the name Lamprothamnus alopecuroides, has proved to be correct, inasmuch as no number as large as 50 had been proposed for any charophyte up to that time. In Europe several forms of Lamprothamnium papulosum (cf. Wood and Imahori, 1965) remain unstudied cytologically. Knowledge of their chromosome number would help clarify their interrelationships. The combination Lamprothamnium macropogon (Braun) Ophel is used here for the plant from South Australia (Wood, 60-9-22-1), with disjunct gametangia and 56 chromosomes. Wood (Wood, 1962a; Wood and Imahori, 1965) designates this taxon as f. macropogon (A. Br.) Wood of var. papulosum of the ‘polymorphic species’ Lamprothamnium papulosum. The value of disjunct 120 CHROMOSOME NUMBERS IN LAMPROTHAMNIUM gametangia as a taxonomic character has been questioned by Wood, but not disproved. The common chromosome number may seem to favour Wood’s treatment, but morphological distinctions in the stipulodes and especially in the disjunct (sejoined) gametangia tend to delimit L. macropogon as a separate species. On the other hand, the monoecious Lamprothamnium with conjoined gametangia (Wood, 60-10-1-5) from Western Australia, previously reported as Lamprothamnium macropogon (Hotchkiss, 1963), is designated here simply as Lamprothamnium sp. The conjoined gametangia preclude referring this form to L. macropogon, while the difference in chromosome number (28 vs. 56) and wide geographical separation are cause for hesitation in assigning it to L. papulosum in the absence of abundant study material. From the data available thus far, the entirely ecorticate, haplostephanous genus Lamprothamnium in its cytology appears to be quite far removed from the morphologically similar section of the genus Chara, the Haplostephanae ecorticatae. It may be noted that the high level of polyploidy in Lamprothamnium contrasts sharply with that in Chara, Haplostephanae ecorticatae, which is almost uniformly at the 14 chromosome level. An exception to this generali- zation is the monoecious species, Chara corallina, reported as n=42 by Sarma and Khan (1965). Chara corallina, however, appears to be a special case in the Haplostephanae ecorticatae, and to bear little relation to the polyploid series in Lamprothamnium which begins at the 28 chromosome level. Another distinction may be seen in the lower (14, rarely 28) polyploid levels found in species with disjunct gametangia (and also in dioecious species) throughout the genus Chara which contrasts with the higher polyploid levels of the game- tangially disjunct species of Lamprothamnium. Cytological data can be correlated with other factors in a preliminary evaluation of the geographical distribution of Lamprothamnium. The wide range of chromosome number found in the Australasian region exceeds that reported from elsewhere in the world and includes the lowest number now known in the genus. Lamprothamnium macropogon is Australian, L. succinctum extends along a well known are of plant distribution ranging from Australasia through the Indian Ocean to Africa. These considerations, together with a considerable range of morphological variation in the Australasian region, suggest the hypothesis that this area is a centre of distribution for the genus. Acknowledgements IT am grateful to R. D. Wood, University of Rhode Island, and to T. Christensen, Universitetets Institut for Sporeplanter, Copenhagen, for the specimens of Lamprothamnium used in this study ; to my wife, Doreen Maxwell Hotchkiss, for continued assistance in counting chromosomes ; and to P. Silva, University of California, for valuable suggestions for the manuscript of this paper. References Groves, J., 1916.—On the name Lamprothamnus Braun. Jour. Bot., 54: 336-337. Hotcuxiss, A. T., 1963.—A first report of chromosome number in the genus Lychnothamnus (Rupr.) Leonh. and comparisons with the other charophyte genera. Proc. Linn. Soc. N.S.W., 88: 368-372. , 1965.—Chromosome numbers in Characeae from the South Pacific. Pacific Sct., 19: 31-37. LINDENBEIN, W., 1927.—Beitrag zur Cytologie der Charales. Planta, 4: 436-466. OpHeL, I. L., 1947.—Notes on the genera Lychnothamnus and Lamprothamnium (Characeae). Trans. roy. Soc. South Australia, 71: 318-323. Sarma, Y. 8. R. K., and Kuan, M., 1965.—Chromosome numbers in some Indian species of Chara. Phycologia, 4: 173-176. Woop, R. D., 1962a.—New combinations and taxa in the revision of Characeae. Taxon, 11: 7-25. , 19626.—Preliminary report on Characeae of Australia and the South Pacific. Bull. Torrey Bot. Club, 89: 250-2538. , and Imanori, K., 1965.—A Revision of the Characeae, First Part. Monograph of the Characeae: 1-904. J. Cramer, Weinheim. MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS N. V. DOBROTWORSKY Department of Zoology, University of Melbourne (Plates iv-v) {Read 29 June, 1966) Synopsis Twenty-nine species have been recorded from Tasmania, twenty-two from Flinders Island, ten from King Island and four from other small islands of Bass Strait. A new species, Anopheles tasmaniensis, and the life history of Culiseta weindorfert (Edw.) are described. The Tasmanian mosquito fauna is closely related to that of southern Victoria. All Tasmanian species either occur in Victoria or have close relatives there. The endemic Culiseta weindorfert is closely related to C. otwayensis Dobr., and Aedes cunabulanus Edw. belongs to a group of the subgenus Ochlerotatus which is well represented in Victoria. Two main elements are represented in the Tasmanian fauna: the southern-Bassian and the northern. The Bassian element is represented by eleven species of Ochlerotatus, three species of Culiseta and one of the apiculis complex of Neoculex. The Bassian element may have originated in the tropics and subsequently spread to the northern and the southern temperate regions, being progressively displaced further from the tropics by later evolving groups. The northern element is represented in Tasmania by fourteen species. Species well adapted to cool conditions are widespread but others are confined to the warmer coastal areas or to warmer habitats. The majority of Tasmanian species are forest mosquitoes and since these have only a restricted ability to disperse across open spaces, it seems that Tasmania received most of the mosquito fauna before separation from the mainland. However, the species which breed in the coastal regions or in the open country may have dispersed to Tasmania, after separation, by the chain of the Bass Strait islands. The distribution of the two forms of Ae. andersoni Edw. suggests that the Grampians form dispersed from the mainland into Tasmania where the typical form evolved and subsequently migrated to the mainland. The main differences of the Flinders Island fauna from that of Tasmania are (1) the absence from Flinders Island of the two endemic Tasmanian species and also those species which are restricted to special habitats not available on Flinders Island, and (2) the presence of two northern species, Ae. macmillani Marks and M. linealis Skuse which do not occur in Tasmania, and of two species which are restricted to the coastal regions and the islands with a mild climate (Ae. clelandi Taylor and Ae. stricklandi Kdw.). The small tauna of King Island reflects the lack of variety of habitats. INTRODUCTION Erichson described the first mosquito from Tasmania, Aedes australis, in 1842. The description was based on two males and one female sent to him from Van Diemen’s Land by A. Schayer. This was followed by Macquart’s description of Aedes nigrithorax in 1847 and Aedes rubrithorax in 1850. Walker described Anopheles annulipes and Aedes crucians in 1856, but the latter is conspecific with Aedes australis Erichson. In 1911 Strickland described six species which he found in collections sent to F. V. Theobald by T. L. Bancroft. However, four of these had been already described and only Tripteroides tasmaniensis and Aedes andersoni (tasmaniensis Strickland) were new species. Taylor described Culiseta littlerti in 1914 and Edwards a further four species in 1926: Culiseta weindorferi, Aedes cunabulanus, Aedes luteifemur and Aedes purpuriventris. Thus during the period 1842-1926 fourteen species were described or recorded from Tasmania. There were no new records until 1948 when D. Lee recorded two more species. PROCEEDINGS OF THE LINNEAN Society or New SoutH Wats, Vol. 91, Part 2 BE 122 MOSQUITOES OF TASMANIA AND BASS STRATT ISLANDS Since 1951 collections from Tasmania and Bass Strait Islands have been made by officers of the Wildlife Survey Section of C.S.I.R.O. (J. H. Calaby, A. L. Dyce, D. L. McIntosh, M. D. Murray and F. N. Ratcliffe) and other entomologists (E. N. Marks, I. Rowley, T. E. Woodward and E. G. Connah). These collections added four species and two subspecies to the Tasmanian fauna (Dobrotworsky, 1953, 1954, 1957 and 1960). By 1960 a total of twenty species and two subspecies had been recorded from Tasmania, four from Flinders Island and two from King Island. This contribution also incorporated the results of collecting trips by the author to Tasmania (October-November 1961, November—December 1962 and October 1963), to Flinders Island (November 1962 and February 1963) and King Island (November 1963). During these studies 255 breeding places were investigated and larvae collected ; almost 2500 adults were collected or reared. In addition to this material I was able to study mosquitoes collected in Tasmania by Dr. M. Littlejohn, and on Bass Strait Islands by Dr. J. A. Thomson, Messrs. S. Murray-Smith and H. Zoski. The island of Tasmania is largely covered with hills and mountains which are particularly rugged in the western part. Forests cover 47% of the island. The temperate marine climate with rainfall throughout the year creates excellent conditions for the breeding of forest mosquitoes. However, the eastern parts of the island are drier with an average yearly rainfall of below 40 inches, and most of the rain water pools usually become dry during November. The western region of the island has an annual rainfall ranging from 50 to 150 inches, and rain water pools do not dry out throughout the year and provide all year round breeding sites for mosquitoes. However, at higher altitudes the develop- ment of larvae would cease during winter where snow may lie for some months. The low-lying areas are restricted and mostly confined to the north and north-east ; they are covered with sclerophyll forest, heath, marshes or tea-tree swamps. Open swamps are rare. Here again in shaded swamps the forest mosquitoes prevail. A description of the physical features, climate and vegetation is given in Atlas of Tasmania 1965. The breeding sites of mosquitoes on Flinders Island are less variable than in Tasmania because the mountains are not high and the island does not experience the cold winter conditions such as exist in Tasmania. Frosts are rare and no snow falls occur even on the highest mountains (the highest Strzelecki Peaks are 2,550 feet). The rainfall does not exceed 35 inches per annum, approximately as high as in the eastern part of Tasmania. The extensive plains with shallow lagoons and swamps provide more habitats for mosquitoes breeding in plain country than in Tasmania. The major features of King Island are the plateau country, the plains, the swamps and lagoons, and dune formations. The highest hills reach only 600 feet. The island provides a very limited variety of habitats for mosquitoes, and only ten species are recorded there as against twenty-two for Flinders Island. A full description of Flinders Island is given by Dimmock (1957) and by Guiler et al. (1958), and of King Island by Stephens and Hosking (1932) and by Jennings (1959). There is a brief review, of both islands, by Littlejohn and Martin (1965). DISTRIBUTION AND COMPOSITION OF THE Mosquito FAUNA Five genera of mosquitoes are represented in Tasmania: (1) Anopheles— four species, (2) T'ripteroides—two species, (3) Aedes—seventeen species, (4) Culiseta—three species and (5) Culev—three species (Table 3). The genus Mansonia is not recorded from Tasmania but is represented by one species on Flinders Island. Their distribution and abundance are determined by various N. V. DOBROTWORSKY 123 environmental factors but particularly by the availability of suitable breeding sites. As would be expected, species which breed in open swamps e.g. Culex globocoxitus, C.p. australicus and An. annulipes are not numerous in Tasmania ; they are restricted to low land areas. In contrast to these, species which are associated with forests are numerous and widely distributed. However, forest mosquito species with special requirements are not evenly distributed: Ae. nigrithorax and Ae. silvestris breed in temporary ground pools in sparse sclerophyll forest; Tripteroides and Ae. notoscriptus are tree hole breeders. TaBLe 1 Distribution according to habitat of larvae in Tasmania Number of breeding sites Ground pools Swamps Species Dis- Total More Com- Tea Rock carded or less pletely Grassy tree pools tyres exposed shaded and to sun tins An. stigmaticus 1 1 An. annulipes 2 1 1 An. tasmaniensis . . 2 2 T. tasmaniensis 2 2g Ae. (O.) migrithorax 4 3 1 Ae. (O.) flavifrons 4 3 1 Ae. (O.) calcariae ee 1 1 Ae. (O.) purpuriventris .. 3 1 D) Ae. (O.) luteifemur 17 7 6 2 2 Ae. (O.) silvestris 6 4 2 Ae. (O.) nivalis .. iil 5 5 1 Ae. (O.) cunabulanus 35 15 i7 3 Ae. (O.) andersont 37 18 15 2 2 Ae. (O.) continentalis 5 3 2 Ae. (F.) alboannulatus .. 9 7 9 Ae. (F.) rubrithorax ae AT 13 31 1 1 1 Ae. (Ff.) rupestris ey 5° 4 2 2 Ae. (P.) postspiraculosis as 2 ) Ae. (H.) australis ae ae 1 1 C. inconspicua. . ie ba 31 6 20 1 4 C. weindorfert .. a ae 2 2 C. ferguson a a ne PG 7 15 4 C. globocoxitus .. Ss ae 2 2 C. p. australicus .. oo She 1 1 255 88 129 12 19 4 3 Species with a greater capacity for adaptation to cold conditions are able to penetrate to higher altitudes where they breed in moorland and sedgeland country ; their larvae are mostly confined there to shallow pools exposed to the sun. These high-altitude species are Ae. nivalis, Ae. cunabulanus and Ae. andersont (Table 2). However, for many species which breed in cool ground water, the range of breeding sites is greater than in Victoria. Thus, species such as C. fergusoni, Ae. rubrithorax and C. inconspicua are not restricted to completely shaded pools ; because of the lower temperatures prevailing during spring and early Summer in Tasmania they are also able to breed freely in pools more or less exposed to the sun (Table 1). 124 TABLE 2 Distribution of mosquitoes according to altitudes MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS Aedes ‘3 shales Ss) SS “-S ~ % i) = = = ages : ae iis odes Localities fs -< Ss s SSS Sees oc | = : 5 & S S z S g S Ss S % ES iy Bo SF Sl Eu Si Gn S US ewe PEP SOAS) G82 AN) RSC OES Vise cms es SS: PSeeese ESS eSobice ESS Piece Onl Om Bridport .10 Sid (ile Maolnuacasc ee cls SS al ae ee wm le ele Sassafras .50 — + | ees ey Sey I) Birralee .50 | a Lake Leake 2) a oe a ft Bae tf a at tL Ee SS Storey’s Creek . 3:000 — — 4 aE ce ee Breona . 3400 + 4+ — + ei DAA YUE RE ee TABLE 3 Distribution of mosquitoes in southern Victoria, Tasmania and Bass Strait Islands Victoria Bass Strait Islands Hast Hastern Otway— Erith, Tas- Species Gippsland Highlands Portland Deal, King Flinders mania and West area, Great Gippsland Dog ANOPHELINAE Anopheles stigmaticus ++ + = aL =n pseudostigmaticus — a at oe) iat Pi atratupes ar == + = == ai ae tasmaniensis . . + — a —_ = ae aL annulipes + + ae = = + al CULICINAE Tripteroides atrupes (Southern form) .. — = =: — == os at tasmaniensis . . ++ + ab — = ah a markst + — = = a iS xaat! Mansonia linealis =F + + — = aL an aurata + — = —_ = ata exe variegata =e — — — = — Me Aedeomyia venustipes + 4 — —_ Zé AM th) Aedes procax ar — — = af ht, ah) theobaldi + — = = eas wo tf al nigrithorax ++ =e at — ait sal amperfectus —- + — — =- —_ ae flavifrons ar =F =F =- + ae ale calcariae + Se au, peed ats td ain N. V. DOBROTWORSKY 125 TABLE 3—continued Distribution of mosquitoes in southern Victoria, Tasmana and Bass Strait Islands Victoria Bass Strait Islands Species Eastern Erith, Tas- East Highlands Otway— Deal, King Flinders mania, Gippsland and West Portland Great Gippsland area Dog purpurwentris clelanda perkinst lutetfemur silvestris nivalis ft camptorhynchus cunabulanus .. andersont continentalis .. aS + aL stricklandi dobrotworskyt notoscriptus alboannulatus rubrithorax rupestris tubbutiensis subbasalis milsont Ae at, + = subauridorsum see + = |< Se PSRSeae hel este IL ae ai cll (= Sloan laeal eg macmillant + multiplex os ok postspiraculosis if + a australis Culiseta victoriensis — + drummondi — + sylvanensis — + hilla is oy = os french ats aa JE + — = = hs = ae ai. =F —- at aL ats eld tet ees |e Wei ele +4] ) 1) ) | | ++++++4+4+1]4144+1 14 ee | otwayensis anconspicua weindorferr litilera . . antupodea Culex fergusont douglast postspiraculosus pseudomelanoconia orbostiensis annulirostris . . globocoxitus we pipiens australicus .. pipiens molestus b+ +++++44 Sipe el | | ae | iN ~J iN oo (SU) pu WS 10 22 30 Although most species are well adapted to cool climatic conditions and non-seasonal rainfall, some are distributed in areas with a warmer climate or are confined to habitats with a warmer microclimate e.g. exposed rock pools. A comparison of the fauna of Tasmania and Bass Srait Islands with that of southern Victoria shows that all Tasmanian species either occur in Victoria or have a close relationship to Victorian species (Table 3). A striking fact emerging from Table 3 is that the mosquito fauna of Tasmania is almost identical with that of the Otway-Portland area of Victoria. 126 MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS In Tasmania, as in Victoria, the mosquito fauna comprises two major elements, (1) the more ancient southern or Bassian element consisting of the genus Culiseta, one group of the subgenus Ochlerotatus and the subgenus Neoculex (apicalis complex) and (2) the northern which is a part of the Indo-Malayan element. This element, which includes a great assemblage of relatively recent successful insects, is represented by Anopheles, Tripteroides, Mansonia, Culex and subgenera Finlaya and Pseudoscusea, and on Flinders Island Chaetocruiomyia, of the genus Aedes. Previously (Dobrotworsky, 1965) the concept of a southern entry via an Antarctic landmass of the Bassian element in the Australian mosquito fauna was rejected. The detailed study of Tasmanian mosquitoes has not led to any modification of the view that this element entered Australia from the north and was progressively displaced away from the tropics by later-evolving elements. The migration route of mosquitoes in eastern Australia was probably along the coast and ranges, a route which for the migration of plants has been extremely important and of great antiquity (Burbidge, 1960). The successful southern dispersal of mosquitoes on the mainland has depended on climate tolerance and adaptability. Some species of Ochlerotatus e.g. Ae. procax (Skuse) have penetrated just across the Victorian border into East Gippsland, Ae. perkinsi Marks has dispersed throughout Hast Gippsland, while Ae. imperfectus Dobr. has penetrated into the Western Highlands as far as Beaufort. Such pattern of migration can also be observed among mosquitoes of the northern or Indo-Malayan element. Dispersal to Tasmania raises other problems. Tasmania was joined to Australia, and Port Phillip Bay and Bass Strait were formerly portions of a continuous land surface (Keble, 1946). A land connection between Australia and Tasmania remained until late Tertiary times; Tillyard (1926) believes that the land connection was broken and joined several times. This land bridge has been regarded as the main route for the migration of flora and fauna Paramonow (1959), however, has questioned the importance of this land bridge. ‘‘ The ability of an insect to fly or to be carried by wind for very long distances, the presence of a group of islands between Tasmania and Australia forming a bridge between them, are factors nullifying the presence of the strait between Tasmania and Australia.’”’ It seems to me that this cannot be applied to all winged insects in spite of many reports of the ability for long distance flights of many of them and the importance of wind in their dispersal. For example, in Canada the adults of tent caterpillar moth (Malacosma disstriata Hubner) were carried by the turbulent air associated with cold front for a distance of at least 300 miles in 12 hours (Brown, 1965). Trapping of airborne insects from ships and airplanes has yielded large numbers of insects (Glick, 1939 ; Gressitt et al., 1960, 1961 ; Yashimoto and Gressitt, 1960, 1961 ; Harrell and Yashimoto, 1963; Madison, 1964). The mosquitoes were collected only on few occasions, but there are reports of their being taken at altitudes up to 5,000 feet, and at sea as far as 110 kilometres from nearest land. It should be kept in mind that the occurrence of active or passive dispersal of insects depends largely on their behaviour and ecology. Insects living in open country or in the upper strata of the forest may be easily carried by winds or may be able to fly long distances. Insects of this type include the southern Victorian mosquitoes: Ae. camptorhynchus, Ae. australis, Ae. notoscriptus, Ae. macmillani, Tripteroides, An. annulipes, C. globocovitus and C.p. australicus. These species are potentially able to cross Bass Strait particularly by “‘ jumping ”’ from island to island. The ability of some of these species to breed in salt or brackish water is a further favourable factor helping dispersal across sea barriers. However, the majority of Tasmanian and southern Victorian mosquitoes are ecologically confined to forests and even during swarming they do not fly in N. V. DOBROTWORSKY 127 the upper strata. Such mosquitoes usually do not fly out of the forests where they have their breeding sites and are not exposed to strong winds which could carry them for long distances and across water gaps. The unfavourable ecological conditions associated with open and dry land constitute another barrier to the dispersal of many forest mosquitoes. Several common species We Pel * KENT GROUP MING ISLAND STATUTE MILES STATUTE MILES ISOHYETS IN INCHES “60~ INDIAN TASMAN 60-100 Inches (Estimated) as 4 Ae. cunabulanus e Ae. nigrithorax ae C. weindorferi 138° 148° Fig. 1. Distribution of some species of mosquitoes in Tasmania and Bass Strait Islands. of Culiseta widely distributed in the Eastern Highlands of Victoria have not dispersed across the intervening plain into the ecologically favourable Grampians and Otway Ranges. Quite short stretches of ocean have apparently prevented the dispersal of the common Tasmanian species Ae. cunabulanus through the chain of islands to the ecologically suitable Flinders Island, itself only some 35 miles off the Tasmanian coast (Fig. 1). Ae. cunabulanus presumably evolved in Tasmania 128 MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS before the Pleistocene but did not cross the land bridge which joined Tasmania to the mainland during the glacial period. It is difficult to imagine the cause of this failure but it is possible that this species evolved in remote or secluded parts of Tasmania and did not disperse until the disappearance of the land bridge. While Bass Strait would seem to constitute an impassable barrier to the spread of some mosquitoes, the distribution of others seems to confirm Paramonov’s (1959) claim that the main obstacles to the spread of some Victorian insects to Tasmania are climatic and ecological. This may be the explanation of the absence from Tasmania of Victorian species such as Ae. dobrotworskyt Marks which occurs in southern Victoria from Genoa to Apollo Bay, and M. linealis and Ae. macmillant which have reached Flinders Island but are apparently absent from Tasmania. The absence of C. weindorferi from the Bass Strait Islands may also be due to climatic factors (Fig. 2). NEW SOUTH WALES SOUTH AUSTRALIA \ King Is. ) 4 Ae. macmillani TASMANIA e Ae. continentalis s An. tasmaniensis \|{\I\\|| am. dinealis Fig. 2. Distribution of some species of mosquitoes in Victoria, Bass Strait Islands and Tasmania. It seems therefore that the main migration route was from Victoria to Tasmania. The distribution of Ae. andersoni suggests however, that the migration of this species has occurred in both directions. The Grampians form of Ae. andersoni is common in the Grampians and is recorded also from a few localities along the Dividing Range to Kiandra in New South Wales. It is likely that this form migrated ‘to Tasmania where the typical form evolved. . In Tasmania this form is highly adapted to cold conditions and often breeds at high altitudes (3,000-3,500 feet). On the mainland its distribution is coastal. The most important differences between the mosquito fauna of Tasmania and the Bass Strait Islands and that of southern Victoria are: (1) absence from Tasmania of the group of northern species which in Victoria are almost entirely confined to Hast Gippsland. (2) The absence from Tasmania of Culiseta Species breeding in subterranean waiters in tunnels of land crayfishes. These Species are common throughout Gippsland. (3) The occurrence in Tasmania N. V. DOBROTWORSKY 129 of two endemic species: Culiseta weindorferi and Aedes cunabulanus. (A) Distribution of Aedes andersoni in Tasmanian highlands up to 3,500 feet. (5) The mosquito fauna of Bass Strait Islands is impoverished Victorian fauna. Paramonoy (1959) described the Tasmanian insect fauna as impoverished by comparison with that of Victoria and attributed this condition to the elimination of many species during the Pleistocene glaciation. However, it would seem that the impact of glaciation on the Tasmanian flora and fauna has been exaggerated, for Jennings and Banks (1958) assert that ‘‘ an ice sheet occurred only in the Central Plateau with cirque and walley glaciers elsewhere ”’. There is no doubt that certain endemic genera of plants became extinct during the most severe phase of glaciation (Burbidge, 1960) and it is possible that some endemic species of mosquitoes suffered the same fate. However, there is no evidence that the Tasmanian mosquito fauna was ever substantially richer than at the present time. TAXONOMIC ACCOUNT Family CULICIDAE Key to genera Adults ih, Scutellum trilobed, each lobe bearing bristles. Palps in female much SHGRCCK: LAMM PLOMOSCIS! “sat t-kts.. Peers Mates ehte Rabo QIAN inh deta a CoAeha ie ots Apo aes totam 2 Scutellum evenly rounded with marginal setae along it. Palps of FEIMAleS MASH OMGHAS! PLOW OSCISM psa: -acaeMe) oto alsieiie ites apleyes a ogo) ope Mel Oe oie sues ou pn den Anopheles eee CS) EM ANE OLESCIGS othe ince ets ce ease © waite Sys cio hea rauere Sater em ee ea eiere See Culex lla lo Seta mics cysrye beled peer ee er Aeaeiats. mikes sabe eisaeans ene Reng ere his Sie wana mlarale Sime e cle 3 San (2) eS pikacular pristles, ab sem bis tee crgec ry. Sipcy. dere cyctoy yore 4c aan bey Raye bee Go cea cua eye ete oe 4 Sikacull Mb cIstlesgOLeSeMUNupr usr ce Gael tre hk) nlag Gian teen aps solutes, cere ecuene Giinegememetee ae 5 fas) Peoctspiracularabristlesi presenti wseeridee = cio ols sical; Setacbeeiedae tiene) - chevelle Aedes Postspiracular bristles absent ......... PPS scales ani SY gs SERIE Phe Pyayeayns Mansonia 5. (3) Uniformly brown; base of subcostal vein hairy beneath ................ Culiseta Black with patches of broad white scales on thorax and abdomen ; base) of subcostal vem bare’ beneath 2!.....3-5.-----5-50+52 esses Tripteroides Larval (fourth stage) I VIIIth abdominal segment without respiratory siphon .................. Anopheles VIlIth abdominal segment with elongate respiratory siphon .................... 2 2. (1) Distal half of siphon modified for piercing roots of aquatic plants ........ Mansonia SHIM MOb maochiticcl momangil iN SAYS coscosncocodococsovscosccsoocsbocsdaoooc 3 3. (2) Anal segment with ventral brush (seta 4) reduced to one pair of SCto Cit tehas oy ead tater ties sata spies sabes oi tks tafe fo anne by tage as Gnptabiapenogele ke oat Trupteroides Anal segment with ventral brush of several tufts .............................. 4 4b, (@)) Sijolaorm, Watda joenie it. joel Pewee coco aeccdcoensobaccedsudadooobe Culiseta (in part) Siphonswithouts basaltsetae& Sit. 9 Sata) SER, BS aes. ata Stele Sede mente « 5 9 (4) Siphon swath) several pairs off ventral tufts! 925505. .se eee es rele ee cee Culex Siphon with a pair of median tufts or setae ..................2..25-55--25-025 6 6. (5) Some of setae plumose; anal segment usually not completely ringed loxyanisaclollow - AICE SARL AAS, AREA EASES cee Aah SNA Rag, AEE 20 TATE RIE De csr Va dae Aedes All setae non-plumose ; anal segment completely ringed by saddle .. Culiseta (in part) Subfamily ANOPHELINAE Genus ANOPHELES Meigen Key to Tasmanian species of the genus ANOPHELES Adults Ile Wings and legs profusely marked with white scales ...................... annulipes Legs dark scaled 2 2. Wings with some silvery or pale sections of veins. Hind femur entirely adarkwscalediyr.:.consayess )..: Sathae § ocsisepl oh A Ge in J EIS eee ch REE potent 3 Wings entirely dark scaled. Basal four-fifths of hind femur creamy SERINE |: Nasco cde ertecn bate acer Honan Game PORCNO eR RRO ance BENE res ice ine Ica stigmaticus 3. Fringe of wings with distinct white section at tip ........................ atratipes Nae OF YynMas EimAlhy Chi | Co oodbanomeaocuouec oso ope ouuc hhibuoodos tasmaniensis 130 MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS Subgenus ANOPHELES Meigen ANOPHELES (ANOPHELES) STIGMATICUS Skuse Anopheles stigmaticus Skuse, 1889, Proc. Linn. Soc. N.S.W., 3: 1758. This is a small brown species. The basal four-fifths of the hind femora are clothed with creamy scales. It has been recorded from Hobart (Dobrot- worsky, 1957). Additional record is: Flinders Island. It also occurs in the mountainous areas of Victoria and New South Wales. Material examined: 24. ANOPHELES (ANOPHELES) ATRATIPES Skuse Anopheles atratipes Skuse, 1889, Proc. Linn. Soc. N.S.W., 3: 1755. This also is a dark brown mosquito. However, the hind legs are entirely dark and the wings have white sections along some veins; the fringe scales at the tip of the wing are white. A single upper sternopleural bristle. The palpi are black and the two basal segments have a rather shaggy appearance (Plate 1). It is a coastal species not previously recorded from Tasmania or Bass Strait Islands. It has been collected by A. L. Dyce and M. D. Murray at Bridgewater, and by the author at St. Helens and Detention River in Tasmania, and on King and Flinders Islands. It also occurs in all mainland states. Material examined: 14, 149. ANOPHELES (ANOPHELES) TASMANIENSIS 0. Sp. Types: The holotype male and three paratype females were bred from larvae collected 29/11/62 at Birralee, Tasmania, and have their associated larval and pupal skins; the allotype and one paratype female were collected at the same locality and date, and eight paratype females were also collected there (27/10/61). The holotype male, allotype female and seven paratype females are in the National Insect Collection, O.S.I.R.O., Canberra. One female paratype is in each of the following collections: National Museum, Melbourne; School of Public Health and Tropical Medicine, Sydney; University of Queensland, Brisbane ; British Museum (Natural History), London; U.S. National Museum, Washington. Distinctive Characters. Adult: A dark species which has pale scales along only some sections of the wing veins. It can be distinguished from An. atratipes by the narrower squama scales of the veins, the entirely dark fringe of the wing, the presence of 3-5 upper sternopleural bristles. The two basal segments of the palpi are not shaggy (Plate 1). Larva: Brown with white dorsal longitudinal stripe on the thorax and abdomen. Holotype Male. Head: Vertex with pale upright forked scales, becoming black laterally. Frontal tuft white. Tori dark, flagellar segments of antennae pale, but two terminal segments dark; verticillate hairs silver grey. Palps dark scaled; long hairs on segments IV and V, pale. Proboscis black. Thorax: Scutum brown, margins lighter; anterior margin with median and lateral white scale-tufts. Bristles dark. One strong and three weaker upper sternopleural bristles and four lower sternopleural bristles; seven-eight upper mesepimeral bristles. Numerous pale setae in front of wing roots and along lateral edge of scutum. Scutellum light brown with 16 long black setae, and several small pale ones. Legs black, hind femora paler ventrally. Wings: Fringe dark without white section at tip of wings (wing scales partly rubbed, See description of wing venation in allotype female). Knob of haltere brown scaled, with pale stem. Abdomen: Dark brown, almost black, clothed with N. V. DOBROTWORSKY 131 numerous pale hairs; scales absent. Terminalia (Fig. 3 a, b): Coxite short, blunt, tapering, with long setae and a few scales laterally ; inner side of coxite with a long subapical seta curved at its tip. A single stout parabasal spine curved at its tip, arising from an elongate base; ventral lobe of harpago with two short and two longer setae; dorsal lobe with fine broadened setae, four of them in close set row, and one shorter seta arising distally between them and ventral lobe. Style long and slender with minute setae along it; terminal appendage small. Lobes of tergite widely separated, with numerous short setae. Fig. 3. Anopheles tasmaniensis n.sp., a-b, adult: a, harpago; 6, phallosome; c-g, larva: c, prothoracic setae 1-3; d, pro-, meso- and metathoracic pleural setae; e, head, mentum and terminal segments; /f, antenna; g, pecten; h-j, pupa: h, abdomen; 7, cephalothorax and metanotum; 7, trumpet. Allotype Female. This differs from the holotype as follows: Tori and first flagellar segment pale, other flagellar segments black. Palps as long as proboscis without labella, black scaled. Scutum dark brown between dorso-central bristles ; fossa and lateral area light brown sparsely clothed with small narrow curved pale scales; bristles black, but pale in front of wing roots and along border with pleura. Scutellum with a row of 25 strong black and several smaller pale bristles. 5-6 lower sternopleural bristles; 3 bristles on prealar knob ; 10-14 upper mesepimeral bristles. Wings: C, Se, R,, R,, Rs and Ms,,, uniformly clothed with dark scales; apical three-quarters of Rs, pale scaled ; R,,; with some pale scales, particularly distally; Cu, pale scaled on basal three-quarters, dark apically ; M and An, pale scaled, some black scales basally. Patches of black scales at base of fork R, and R;, and M, and M,, at base R,,;, and M—Cu where it joins Cu. 132 MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS Paratype Females. The series of 12 paratypes do not show significant variations. Larva (Fig. 3 e-g): Brown with white dorsal longitudinal stripe on thorax and abdomen. Head: setae 2, arising close together, long, stout, single and simple; 3, single, simple; 4, fine, single, simple; 5, 6 and 7 plumose almost equal in length; 8, small, about 4 branched; 9, larger, about 7 branched. Antennae spiculated ; seta 1, arising at about mid-length, about three-quarters as long as antenna, plumose. Thorax: Prothoracic setae 1 and 3, single, simple; 2, about 6 branched. Metathoracic palmate setae (1) reduced to branched hair. Abdomen: Segments I-VIII with narrow transverse tergal plate. Pecten with about 24 spines. Anal segment: Saddle covered with fine spicules ; seta 1, long, single; 2, about 13 branched ; 3, about 7 branched ; 4 (ventral brush) of 14-16 plumose setae. Pupa (Fig. 3 h, 7): Trumpet broad with almost quadrangle opening, with deep incision. Paddle: without fringe; seta 1, single, stout; seta 2, small, fine, 5-6 branched. Eggs are illustrated in Plate 2. Biology: The adults were common in a tea-tree swamp near Birralee, northern Tasmania 27/10/61, but no larvae were found at that date. On the second occasion (29/11/62), this swamp was very shallow and dry in places ; several larvae and pupae were collected in the deeper parts. Distribution. Tasmania: Birralee and Redpa. It also occurs in a coastal belt of Victoria. Material examined: 1,4, 209. Subgenus CELLIA Theobald ANOPHELES (CELLIA) ANNULIPES Walker* Anopheles annulipes Walker, 1846, Ins. Saund. Dipt., 1: 433. Mackerras (1927), Lee and Woodhill (1944) pointed out the variability of this species in Tasmania as well as on the mainland. Specimens with an entirely black proboscis and others with the proboscis pale on the apical half have been collected in Tasmania. It has been recorded from Launceston and Low Head (Lee, 1948). Additional records: Bridgewater (A. L. Dyce and M. D. Murray). Little Swanport, St. Patrick’s River; King and Flinders Islands. It is widely distributed on the mainland of Australia. Material examined: 4,4, 19. Subfamily CULICINAE Tribe SABETHINI Genus TRIPTEROIDES Giles Subgenus RACHTIONOTOMYIA Theobald Atripes Group TRIPTEROIDES (RACHIONOTOMYIA) ATRIPES Skuse Culex atripes Skuse, 1889, Proc. Linn. Soc. N.S.W., 3: 1750. A small black mosquito without a white pattern on the scutum; the pleura are densely clothed with white scales. *For synonymy of this and other species, see Dobrotworsky (1965). N. V. DOBROTWORSKY le This is a first record of this species from Tasmania. All four specimens examined belong to the southern form of YT. atripes which have small lateral patches on the tergites visible from above only on segments VI and VII. The females have been taken biting at Lisdillon and Ferntree. T. atripes also occurs in South Australia, Victoria, New South Wales and Queensland. Material examined: 49. Caledonicus Group TRIPTEROIDES (RACHIONOTOMYIA) TASMANIENSIS (Strickland) Stegomyia tasmaniensis Strickland, 1911, Entom., 44: 249. This can be distinguished from 7. atripes by having a band of white scales across the pleuron, and by the pale scaling of the terminal segments of the hind legs. It is distributed mainly in areas with an annual rainfall higher than 30 inches. This species has been recorded from: Advent Bay, Bridport, Cradle Valley, Devonport, Eaglehawk, Geeveston, Hartz Mt., King River, Launceston, Mole Creek, Mt. Arthur, Mt. Farrel, Mt. Wellington, New River, St. Patrick’s River, Springfield (Edwards, 1924, 1926); Ferntree, Low Head (Lee, 1946, 1948) ; additional records are: Maydena (D. L. Dyce and M. D. Murray); Derwent Bridge, Renison Bell and Flinders Island. It also occurs in Victoria, South Australia and New South Wales. Material examined: 64, 209. Tribe CULICINI Genus MANSoNIA Blanchard Subgenus COQUILLETTIDIA Dyar MANSONIA (COQUILLETTIDIA) LINEALIS (Skuse) Culex linealis Skuse, 1899, Proc. Linn. Soc. N.S.W., 3: 1747. Distinguished by the light-golden longitudinal lines on the dark-bronze sealed seutum and by the white scaled venter with median black patches and black apical border on the sternites. This species has not been found in Tasmania but occurs on Flinders Island (J. H. Calaby and D. L. Dyce, 2/12/51; N. V. Dobrotworsky, 10/2/63). This is the most southern record of M. linealis which occurs in South Australia, Victoria, New South Wales and Queensland. Material examined: 89. Genus AEDES Meigen Key to species of the genus AEDES Adults, female I Marsal scomenbs awallan TOG uvylitem ko ciri Gls) Merereteyete renee ieieiel tele ictetel-ielelaie ysis tay eevee 2 Tarsi darkescaleds wibthoutmwhite bands "Sosseee 120s... weenie scree . . 10 Deny a\Wanesemnovilediwiblimwinibenscales:” srl. fc crs clots eile ene eters siavee a eicnereee ee Wings dark scaled or with a few slightly paler scales .......................... 3. (2) Wings with a few slightly paler scales ........0-..0.5... 55.00... form A Wines Commas Chik senllcl coccscvdcccbons cmos ouauousucco Loon dbcbunagonouGk 4. (3) Scutum with longitudinal lines and lyre-shaped pattern ................ notoscr on. Sen, {MAO HOLY JOC A Sebo ose ooo tooo cSubiod dou bbo boo oo oboc.cb ofl oeoe 5 134 MOSQUITOES OF TASMANIA AND BASS STRAIT ISLANDS 5. (4) Scutum with patches of white scales ; femora with ochreous preapical 10 Um PE rE ial Do PRA NS Aa ares se; ce) RDN MUM ils notes ae Sicha: guchty 6 Bed alboannulatus No patches of white scales on scutum ; femora without preapical rmg .............. 6 65 1() kitth tarsal gsesment) jotpehinclgle sso ac qe ay eee serra eee eee nnn een ne ee 7 BitthytarsalecesmentemonrenOrslessimyjait esc: LOC ile iin aee eee eee eres 8 7. (6) Venter more or less ochreous, usually with scattered black scales ........ rubrithorax Venter either black with white lateral patches on sternites, or white withweapicall blacks bandmonmsternitesm mens er sii keene eee eens rupestris 8. (6) Fifth tarsal segment of hind legs all white ; legs unmottled ; proboscis black: 1seale de ipete:s 5 PLY ‘i ov ae da atin Zaseitie dr nae ods tanner ae uh ree ae A OW? x ae ’ ‘og 1G Lalande ie Tae BuES cat. oe y “hot : aes: Ca ie ie 4 ee: : te pemee) are pee 2 a) ree #(2)) a te ee aaa! Ad "dion oar w in isey an ie ye 6° dra! Lidd Beit at arto - eagieelrn pared GE) SM (a Nit * Te. iat tee inapatiie os ic etme. 6; marae. - egeh ophae teecls i ee! woigaiansion) Pee ime pg oaht ‘ Ba rset Ses Paes iy rise wei del es visthwbieten, YOR! - Aq the mare * ny Proc. Linn. Soc. N.S.W., Vol. 91, Part 2 Anopheles atratipes Skuse and Anopheles tasmaniensis Dobrotworsky. PLATE IV ey Cae Vedi’ glia b yee Ti oie! : : <— Proc. LINN. Soc. N.S.W., Vol. 91, Part 2 PLATE V Anopheles tasmaniensis Dobrotworsky and Aedes nigrithorax Macquart. STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS TV. EXPRESSION OF ALLELOMORPHISM AND GENE INTERACTION FOR CROWN RUST RESISTANCE IN CROSSES INVOLVING THE OAT VARIETY BOND EK. P. BAKER and Y. M. UPADHYAYA Department of Agricultural Botany, The University of Sydney [Read 28th September, 1966] Synopsis Segregation for crown rust reaction type was studied in the F, and F, generations in both seedling and adult plant stages in crosses between the resistant variety Bond and susceptible varieties and in crosses between Bond and the varieties Landhafer, Santa Fe, Trispernia, Mutica Ukraine (Ukraine) and Victoria, all resistant to the prevalent Australian races. The immunity of Bond was conditioned by two dominant genes, acting in complementary fashion, in both stages of growth. However dominance was incomplete at high temperatures, modifying the characteristic F, 9 resistant: 7 susceptible ratio to a corresponding 5: 11 ratio. The studies extended the genes for resistance identified in certain of the resistant varieties. Landhafer was found to have a third factor acting in complementary dominant fashion with one of the two complementary Bond factors in conditioning seedling resistance only. A third factor in Santa Fe was a complementary dominant factor to the same member of one of the two Bond factors. It was effective in conditioning resistance in both seedling and adult plant stages. The fifth factor in Ukraine was a third member of the allelic series at one of the Bond loci. Fulghum and Algerian contributed the fourth member of the allelic series at one of the Bond loci and acted in complementary fashion with the second Bond gene to confer adult plant resistance only. Grass clump type segregates were observed in the F, generation of the cross between Bond and Landhafer. The segregation ratio suggested the genetic action of an inhibitor gene to a dominant gene for grass clumping. INTRODUCTION The oat variety Bond has proved a valuable source of resistance to crown rust (Puccinia coronata avenae Erikss.) in North America in breeding improved varieties. It originated in Australia as a selection from a cross made by Pridham at Cowra Experiment Farm between the variety Golden Rain and an offtype of the Red Algerian variety introduced from Algeria. Resistance to most races of crown rust and some races of oat smuts was incorporated into Bond; resistance to the latter diseases apparently was transferred from the Red Algerian parent. Reference to the mechanisms involved in the incorporation cf crown rust resistance will be considered later. In addition, Bond possesses the agronomic virtue of stiff straw. With the increase in prevalence of crown rust races attacking Victoria derivatives in North America and the susceptibility of these varieties to Helminthosporium blight (Helminthosporium victoriae Meehan and Murphy) considerable attention was turned about 1945 to the production of commercial varieties deriving resistance from Bond. Several important economic varieties were subsequently released from Bond crosses. However, with the popularity of such derivatives, race 45 of crown rust, capable of attacking Bond and its derivatives, increased tremendously. As a result the varieties Landhafer, Santa Fe and Trispernia have been used more recently in breeding for rust resistance. In Australia Bond, included in the international differential varietal crown rust set in physiological race surveys, was resistant until 1949, when Waterhouse (1952) isolated two races capable of attacking it from field collections. Investi- gations by Baker and Upadhyaya (1955) for the period 1953-55 showed that races virulent on Bond were not prevalent and Bond remained a good source of resistance for the breeding of resistant varieties. PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WaAtzEs, Vol. 91, Part 3 156 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV In the results presented in the current paper the mode of inheritance of crown rust resistance in Bond was analysed. The factors present in Bond were also shown to interact, or to be allelic, with factors in other resistant and, in certain instances, susceptible varieties. Genetic independence or otherwise between the Bond genes and the stem rust resistance gene (Rd,) in Richland was also studied. LITERATURE REVIEW Crown rust resistance in Bond has been shown to be conditioned by a pair of dominant complementary factors by several investigators (Hayes et al., 1939; Weetman, 1942; Cochran ef al., 1945; Ko et al., 1946; Litzenberger, 1949; Kehr and Hayes, 1950; Griffiths, 1953). However, Torrie (1939) obtained an F, ratio approximating 11 susceptible : 5 resistant plants, and interpreted the results on the basis of the action of a strong gene in Bond with another masking its effect. At the same time this ratio was explained by Hayes et al. (1939) and Ko et al. (1946) on the basis of a very weak expression for resistance when both complementary factors in Bond were present in the heterozygous condition. Modifications of the above reported ratios were observed by certain workers. Apparent segregation of a single factor due to the common presence of one of the complementary Bond genes was reported in crosses between Bond and the varieties Iogold (Hayes et al., 1939) and Santa Fe (Litzenberger, 1949). Whilst Finkner et al. (1955) reported the operation of a single factor in Bond from observations on the cross of its derivative, Clinton, with Ukraine, Osler and Hayes (1953) detected the presence of non-identical allelomorphic factors in the variety Santa Fe. Weetman (1942) from field data indicated that resistance in Mutica Ukraine to race I was due to two dominant complementary genes. He suggested that the factors in Ukraine were probably allelomorphic to the Bond genes for resistance. Inhibition of the factors for resistance in Bond has been reported by certain workers. Richland-Fulghum possessed two complementary inhibitors (Cochran et al., 1945), Ukraine possessed one factor for seedling resistance to race 57 and this gene inhibited the resistance of Clinton against race 109 (Finkner e¢ al., 1955); undetermined factors in 8.D. 334 (a Richland derivative) inhibited the resistance of Bond resulting in an F, ratio of 15 susceptible: 1 resistant plant (Ko et al., 1946). Studies on the mode of inheritance of crown rust resistance in the varieties Landhafer, Santa Fe, Mutica Ukraine and Victoria have been reviewed in previous papers in this series (Upadhyaya and Baker, 1960; Upadhyaya and Baker, 19626). Against Australian races Landhafer possessed two factors for adult plant resistance, one of which conditioned seedling or physiological resistance in addition ; Santa Fe possessed one factor operative in both seedling and adult plant stages; Mutica Ukraine (henceforth designated as Ukraine) possessed one factor for seedling resistance independent of the three factors for adult plant resistance (two of which acted in complementary fashion). The Victoria type crown rust resistance was conditioned by two linked complementary factors (with a linkage value of 9-6 +1-7) in the seedling stage, by two independent factors operative at the adult plant stage only and by one independent factor operative at both stages. The action of the latter factor was inhibited by a linked gene showing approximately 10°, recombination with it. One of the factors for adult plant resistance was linked with approximately 10% recom- bination with the two linked complementary factors for seedling resistance. Upadhyaya and Baker (1965) studied the relationships of the genes for resistance in the varieties Landhafer, Santa Fe, Trispernia, Ukraine and Victoria to the prevalent Australian races of crown rust. The two factor pairs in Landhafer conditioning adult plant resistance, one of which conferred seedling E. P. BAKER AND Y. M. UPADHYAYA 157 resistance in addition, were independent of the factors in the varieties Santa Fe, Trispernia and Victoria. The seedling reaction type of Landhafer was epistatic to those of Trispernia and Victoria. The factor for both seedling as well as adult plant resistance in Santa Fe was independent of the factors in Victoria and epistatic to them. Certain modifying gene(s) resulted in the expression of a reaction type similar to that characteristic of Victoria by suppressing the Santa Fe gene. The Santa Fe factor was considered allelic to the factors in Ukraine and Trispernia. The reaction type of Santa Fe was dominant to that of Trispernia in tests with four races, but with race 203 the Santa Fe gene was inhibited by a pair of complementary factors, one contributed by each variety. The three factor pairs in Ukraine, two acting in complementary fashion, involved in adult plant resistance were independent of the Santa Fe gene. Indirect evidence indicated that the factors conditioning seedling resistance in Santa Fe and Victoria were genetically independent. The independence of the factors responsible for adult plant resistance in Landhafer and Ukraine and likewise the independence of the Ukraine and Victoria adult plant factors could not be established in the absence of studies involving the appropriate crosses. Upadhyaya and Baker (1962a) showed that the oat varieties Burke and Laggan possessed a single gene (Rd,) operative against races 2 and 12 of stem rust (P. graminis avenae KE. and H.). EXPERIMENTAL RESULTS The experimental procedure was as set out by Upadhyaya and Baker (1960). (A) Studies on Inheritance of Crown Rust Resistance Crosses involving Bond with the following varieties were studied :— Fulghum, Algerian, Burke and Laggan—susceptible to all crown rust races used. Santa Fe, Landhafer, Ukraine and Victoria—susceptible to only specific races of crown rust. (a) F, Reaction Types and Reactions The reaction types and reactions of the parents and F,’s are presented in Table 1. The adult plant immune reaction and highly resistant reaction seedling type (‘‘0;’’) characteristic of Bond was dominant or partially dominant in crosses involving specific races where the second parent was susceptible. Likewise where both parents were resistant the immunity or highly resistant reaction type of Bond was usually epistatic or partially so. However, in certain eases a slightly less resistant reaction than either parent was observed in the mature plant stage. Incomplete dominance of the reaction or reaction type of resistant parents was observed in the case of race 203 to which Bond was susceptible. To field inoculum, comprising a mixture of races, none of which were, however, virulent on Bond the reactions of the F, plants in Bond crosses were intermediate or moderately susceptible. In certain cases slightly variable reactions were exhibited among different F, plants in the same cross. (b) Behaviour of Segregating Generations Involving Races to which only Bond was Resistant (1) F, Segregation (i) Seedling tests Data relating to these tests are presented in Table 2. In crosses between Bond and Fulghum, Algerian, Burke, Santa Fe and Ukraine a satisfactory agreement between observed and expected results for each F, population was observed on the basis of segregation of two dominant complementary factors contributed by Bond. The total segregation of the separate plants in each cross and the grand total for all crosses also agreed STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV 158 ‘e1qijdoosns=g ‘uorjover oyerpoutioqUI="quT “yURAQSISEA Ajoyeiopou= YW “4ueysiser=y ‘ounurut—Tq , yyy W = = I = =[-: ==! <% I ot) ug a puog x Brmojora Ty W—qUL VIN = WIN I W =1‘'0 £0 re £0 £0 I ‘* puog x deyeypuey typ = T = if I = o = 0 = = = ba puog x oureryg Tip WIN ‘I = = W = W a) - og —§ = £0 06 ‘T °° puog xXeq eyueg Tq S—"qU] YIN WIN SAAN I = : a = = og—5 o me puog Xuessey iy y-qUyl = = = = = = - = = =e = a "*puog x eying "yp g—"qUuy, W W = W = £0 £0 - - a) = puog x uewesty Wy UW = UN IN W = I-'0 £0 = - = = “7 puog x umysny Ty (%86) I S W W I WT uy ¢ Uy uy uy Uy ce ae “* BLIOJOLA, I I IT il I I ig ‘ : : : : i ee "+ geyeypuey “qUI-YWN ‘T WIN ii if Il I v P : : 7 P < S "* oureIy.) I iL I T iL Il 2 : ‘ : : : ae zs "* oF Byueg SI = = = = = F F i F F F uesseT Sg = = = = = ¢ 12 F F 12 i Zz ae eying (%86) S = = = = = ¢ if 12 P ig P e ee "* wellosTy Ss Ss Sg g Si Si g Pv i F 7 P “3 3 “* umyYysny I I i I I 8 0 1) a) om) 1) F pe puog 6&6 F-LEG LEG 92% £0Z 982 68z F-LES LES 92Z £0% PIS SSOJO JO JVUuSIe eseqs qnups eu ul SUOTIOBOY sooBel O4 SUOTIOVOL SuI[peeg puog fjarvoa yoo oy; burajoaus sasso1o ur suoyones ynpo pun sadhy uovones buypaas "4 pun joyuasog I @1avy, BE. P. BAKER AND Y. M. UPADHYAYA 159 statistically with the expected 9 resistant: 7 susceptible F, seedling ratio. The data for the totals of each cross were also homogeneous. Figures for the crosses Fulghum x Bond and Burke x Bond, tested against race 286, where the frequencies were corrected from F, behaviour, are not included in the Table. TABLE 2 F, seedling segregation for crown rust reaction type in crosses involving Bond and certain susceptible varieties Crown F, reaction types Cross rust Total Expected ratio P value race Resistant Susceptible 0; 12 3-4 Fulghum x Bond .. 226 Jig) aut a 122 283 9 Res.: 7 Sus. 0:90-0:80 259 60 10 —- 58 128 9 Res.: 7 Sus. 0-80—-0-70 Total 231 180 411 9 Res.: 7 Sus. 1-00 Algerian x Bond .. 226 54 8 — 46 108 9 Res.: 7 Sus. 0-90-0- 80 237-4 30 21 —- 50 101 9 Res.: 7 Sus. 0-30-0-20 286 63 45 2 97 207 9 Res.: 7 Sus. 0:50-0:30 Total 223 193 416 9 Res.: 7 Sus. 0-30-0-20 Burke x Bond 29 89 50 2 125 266 9 Res.: 7 Sus. 0:30-0:20 Santa Fex Bond .. 286 238 80 43 274 635 9 Res.: 7 Sus. 0-90-0-80 Bond x Ukraine .. 226 165) 12) y= 150 327 9 Res.: 7 Sus. 0-50-0-30 Grand Total 1133? 922 2055 9 Res.: 7 Sus. 0-50-0-30 Bond x Victoria .. 259 12 - 16 63 91 5 Res.: 11 Sus.t 1-00-0-90: Landhafer x Bond 286 137 13 - 54 204 45 Res.: 19 Sus. 0-50-0-30 1 Tests conducted at high temperatures modifying expected 9 res.: 7 sus. ratio. 2 16; heterogeneity for totals of 5 separate crosses=1-:96; P value=0-80-0-70. On this calculated basis the number of resistant and susceptible seedlings were 106-1 and 98-9 respectively (P value=0-20-0-10); in the cross Burke x Bond the corresponding numbers were 184-5 and 114-5 respectively (P value= 0:10-0:05). FE, segregation in the cross Bond x Victoria was studied at high temperatures with daily maxima in excess of 80°F. Under these environmental conditions the doubly heterozygous plants exhibited a “3” reaction type resulting in a 5 resistant: 11 susceptible F, ratio with which the observed Segregation agreed statistically. In the cross Landhafer ~ Bond a ratio very close to 3 resistant : 1 susceptible was obtained. It was found, however, from F, studies to be reported later, that an alternative complementary factor from Landhafer interacted with one of the two Bond complementary genes. On this assumption the following genotypes were expected when segregation was studied in tests involving race 286: F, (Resistant) (Susceptible) F, (Homozygous resistant) (Segregating) (Homozygous susceptible), 1 Bd,Bd,BdpBdpBd,Bd, 2 Bd,BdaBdp bd» bd, bd, 1 BdaBda bdp bdp bd, bd, 2 2° 29 ” 2° ” 1 4 2° ” ” 1 TI 2 3” bda ” ” ” ” 1 2° 9° 2° 2” bd, bd, 2 29 ” bdp bdp 2” 2” i & 99 ” ” ” ” 2 4, » BdpbdpBd,Bd, 2 Bd, bd,BdpBdpBd,Bd, Ton oe TENG IB VHB! ENGI: 1 oy) 99 bdp be} be) 2° 4 be) 2° Le) Ly) by) b 1 2 ” 2° ” ” 29 1 2 29 29 29 29 bd, bd, 1 39 Ee) te) 29 bd, bd, 4 ,, ,, BdpbdpBd,Bd, 2 4, 5, Bdp bdpBd,Bd, 8 29 9 2? 29 29 il 4 ty) 29 °° 2” 29 1 4 ”° 99 3° ” bd, bd, 2 ” °° ” ” bd, bd, 2 4, 5, bdpbdpBd,Bd, 1, 5, bdpbdpBd,Bd, 4 29 Le) 99 29 29 1 2 39 99 Le) 29 be) al 7 38 19 =64 160 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV In the F, therefore, a ratio of 45 resistant: 19 susceptible seedlings was expected and a satisfactory fit to this hypothesis was observed. Relationship of F, seedling reaction types to different races was studied in the crosses Fulghum x Bond and Burke x Bond where inoculations on the primary leaves were carried out with mixtures of races 226, 259 and 286 and 226, 237 and 237-4 respectively. In no case was any marked variation in reaction type noted in the case of any F, seedling, indicating the operation of the same factors in Bond against all races in the mixtures. (ii) Adult plant tests. The F, segregation data for adult plant behaviour pertaining to the crosses of Bond with Fulghum, Algerian and Laggan are presented in Table 3. TABLE 3 F, segregation for adult plant crown rust reaction in crosses involving Bond and certain susceptible varieties Adult plant reactions Cross Total Expected ratio P value t R-MR MS-S Fulghum Bond .. ae 65 29 35 129 3 Res.: 1 Sus. 0-70—0-50 Algerian x Bond Bi so ty 39 54 175 3 Res.: 1 Sus. 0-10—0-05 Total ita 215 89 304 3 Res.: 1 Sus. 0-10—-0-05 Laggan x Bond oe .. 126 45 130 301 9 Res.: 7 Sus. 0-90-0-80 1T=immune, R=resistant, MR=moderately resistant, MS=moderately susceptible, S=susceptible. These figures confirmed the segregation of the two dominant complementary Bond factors in the cross Laggan x Bond. However, the data for the other two crosses approximated a 3 resistant: 1 susceptible ratio indicating the action of a non-identical allele to one of the Bond genes in the varieties Algerian and Fulghum, operative in the adult plant stage only. The presence of such a factor in the variety Fulghum was confirmed from studies on the relationship of F, TABLE 4 Adult plant reactions of F', seedlings classified for reaction type to race 286 i crosses of Bond with Fulghum and Burke Seedling reaction types Adult plant reaction Resistant Susceptible Resistant Susceptible 0; s;s—-l 1-—2- 3-4 0; ;-l 1-2- 3-4 Fulghum x Bond Burke x Bond Tt 4 3 - 8 27 = = - R 2 — 1 1 9 15 39 = MR .. See ae 1 2 1 4 - 4 2 Total 15 (17-0)? 10 (9-0) 98 (100-0) 2 (0-0) MS-S 1 1 = 1] (12-0) = = 2 27 (29-0) 2 (0-0) 2 (0-0) 1T=immune, R=resistant, MR=moderately resistant, MS=moderately susceptible, S =susceptible. 2 Expected values in brackets. seedling reaction types and adult plant reactions, data for which are presented in Table 4 relating to crosses of Bond with Fulghum and Burke. Im the cross Fulghum x Bond with the operation of such a factor three out of seven susceptible seedlings would be expected to show resistance at the adult plant stage and all the resistant seedlings would be expected to maintain their resistance as mature E. P. BAKER AND Y. M. UPADHYAYA 161 plants. From the resistant class of seedlings two were found to give a moderately susceptible mature plant reaction and these on progeny tests showed segregation. Obviously these were incorrectly classified. From among the susceptible seedlings, 10 compared with the nine expected out of 21 gave a resistant reaction due to the operation of the allele from Fulghum. In the cross Burke x Bond two resistant seedlings out of 100 exhibited a moderately susceptible adult plant reaction and in this case also these were almost cetainly due to incorrect classification. Similarly two susceptible seedlings produced moderately resistant adult plants, probably for similar reasons. Otherwise there was close agreement between seedling reaction types and adult plant reactions in this cross. It was thus concluded that the varieties Algerian and Fulghum possessed an effective allele to one of the complementary dominant factors in Bond and that Burke and Laggan carried the recessive allele at this locus. (2) KF, Seedling Segregation Progeny tests were carried out on F, plants which had been classified for reaction type or randomly selected. Data for the different crosses are presented in Table 5. The expected reaction in all crosses except Landhafer x Bond TABLE 5 fF, seedling behaviour for crown rust reaction in crosses involving Bond and certain susceptible varieties Crown F, behaviour Cross rust Total Expected ratio P value races Res.t Seg. Sus. Fulghum x Bond 226 259 & 286 @ ea ae 3 22 13 38 1 Res.: 8 Seg.: 7 Sus. 0-50-0-30 (@))... 26 ohn 11 65 64 140 1 Res.: 8 Seg.: 7 Sus. 0-70-0-50 Total 14 87 77 178 1 Res.: 8 Seg.: 7 Sus. 0-70—0-50 Burke x Bond .. 226 237-4 & 237 (a) AF ot 8 57 30 100 1 Res.: 8 Seg.: 7 Sus. 0-30—-0-20 (b) 50 ie 8 70 49 127 1 Res.: 8 Seg.: 7 Sus. 0-50—-0-30 Total 160 «127 84 227 | Res.: 8 Seg.: 7 Sus. 0-20-0-10 Laggan x Bond... 226 NORE SOS 237 I Res.: 8 Seg.: 7 Sus. 0-50-0-30 Santa Fex Bond 286 (b) Ae oe 6 TS | 149 1 Res.: 8 Seg.: 7 Sus. 0-50—-0-30 (c) ae om 12 92 ee 163. 1 Res.: 8 Seg.: 7 Sus. 0-90—-0-80 Total 18 150 144 312 I Res.: 8 Seg.: 7 Sus. 0-70-0-50 Bond x Ukraine 286 (b) 3 ne 7 63° 68 138 1 Res.: 8 Seg.: 7 Sus. 0-50-0-30 Bond x Victoria 259 (d) a Pre 8 453 56 109 I Res.: 8 Seg.: 7 Sus. 0-20—-0-10 Grand Total 73 591 537 42.1201 1 Res.: 8 Seg.: 7 Sus. 0-80-0-70 Landhafer x Bond 286 (c) we ae 12 708 30 112 7 Res.: 38 Seg.: 19 Sus. 0-80-0-70 1 Res. =homozygous resistant, Seg.=segregating, Sus.=homozygous susceptible. 2 (a)=unclassified for F, phenotype, (b)=classified for F, seedling reaction type, (c)=classified for F, adult plant reaction. 3? Some lines in the segregating category showed a preponderance of susceptible plants. was 1 homozygous resistant: 8 segregating: 7 homozygous susceptible lines. In the Landhafer x Bond cross the corresponding expected ratio as shown previously was 7: 38: 19 respectively. In all cases there was satisfactory agreement statistically between observed and expected results. The confirma- tion of the predicted ratio in the Landhafer x Bond cross indicated that Landhafer possessed a dominant factor complementary to one of the two dominant complementary genes in Bond. 162 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV In the crosses of Bond with Landhafer, Santa Fe, Ukraine and Victoria certain segregating F, lines showed a preponderance of susceptible seedlings. This was probably associated with high temperatures and was observed especially in tests conducted under glasshouse temperature regimes between 75°-85°F. F, segregation was also studied in the progenies of F, plants classified for seedling reaction type to race 286 in crosses of Bond with Fulghum and Burke respectively, and these results are presented in Table 6. Good agreement was TABLE 6 F, seedling behaviour for crown rust reaction of I’, plants classified for seedling reaction type in crosses imvolving Bond with Fulghum and Burke F, behaviour to mixture of races F, reaction types to race 286 226, 259 and 286 226, 237 and 237-4 Res. Seg. Sus. Res. Seg. Sus. Fulghum x Bond Burke x Bond 0; ou ae = af rece mn LP 28 1 10 29 — 3-1 = 34 — 1 14 — 1-—2- = 4 — — 39 1 3-4 = 2 59 — 4 29 1 Res.=homozygous resistant, Seg.=segregating, Sus. =homozygous susceptible. shown between F, reaction type and expected F, segregation except for two instances, one in each cross, where a resistant F, plant gave homozygous susceptible progenies. These were probably errors in F, classification. Similarly a few plants classified as susceptible in F, gave segregating F,; progenies. This may have been due to high temperature effects resulting in heterozygous genotypes involving Bond genes being classified as susceptible. (c) Behaviour of Segregating Generations Involving Races to which both Bond and the Second Parent were Resistant (1) F, Segregation (i) Seedling tests. The varieties Landhafer, Santa Fe and Ukraine each possessed a single factor for seedling resistance to races to which they were resistant. The F, segregation in the cross Landhafer x Bond in tests involving race 286 to which Landhafer was susceptible conformed to 45 resistant : 19 susceptible. Therefore with the operation of one additional dominant factor in Landhafer in tests involving races to which it also was resistant the expected F, ratio was 237 resistant: 19 susceptible; if allelism was involved the expected ratio would be modified to a characteristic dihybrid ratio of 15 resistant : 1 susceptible. In the case of crosses between Bond and both Santa Fe and Ukraine the ratio conformed statistically to a 15: 1 rather than 57: 7 ratio. Hence against races to which either Santa Fe or Ukraine were respectively resistant a non- identical allele to one of the two complementary factors in Bond was operative. These were alleles rather than identical genes since the factor in Ukraine conferred resistance to fewer races. The F, segregation of the Victoria type of rust resistance in crosses with susceptible varieties was 71-9 resistant: 28-1 susceptible ; with the operation of the two complementary Bond factors only 12-3 per cent of plants were expected to be susceptible. Since partial dominance was exhibited by the majority of factors in both Bond and Victoria, the intermediate and resistant classes were grouped together for goodness of fit tests. 163 EH. P. BAKER AND Y. M. UPADHYAYA SGP (SS SS) OL:0-02:0 ‘sng 1:8 = ‘seu 6:1L L6P GL 6P IPI GES [849.1 08:0-06:0 ‘Sng 1:86 : ‘Seu 6:1L SIZ 9G GE 19 66 0¢-0-0L:0 ‘sng [:8¢ : ‘SoU 6-1L POL €&@ 9 GG 08 10:0-Z0:0 “Sng 1:86 * ‘Seu 6:1L CII €@ II GS 9G €0€ = ; ; 06:0-00:T ‘sng 4 : ‘seu LG EGE 0 L FOL G61 GL6 = Fi Boe ee GO-0-01-0 ‘sng 4 : ‘soy Lg GZOL Os 9 G66 919 [84O.L 0€-0-0¢-0 SSDS 9L =: (Soul LG G8G GE Tg 006 66Z OL:0-06:0 ‘sng 4 : ‘seu LG OLG Il => LP GIG 0¢:0-0€-0 ‘sng LL :‘sey LG ELI L €T SP SOT 68 eS Se eS SS ee 0€-0-09:0 ‘Sng 61 : ‘Sou LES 696 G9 I 08 €18 1®79,.L 0L-0-08:0 Sng 6 + ‘SeU LES GO ill a LT PLI 08-0-06:0 ‘nS 61 :‘SeuU LES 961 GT T LG ST 0Z:0-0€-0 SnG 61 : ‘SOU LES sol Il == LI 08 OT:0-02:0 ‘Sng 61 ‘rS°8U LES OS? GG —s 61 90P vas —G = =§ >-!0 efqtydoosng queqsisoy onjea q oer poyoodxa [eO0, sedéq4 uomovor * 7 ‘eTqiydoosns = ‘sng 986 V-LESG 9%Z 0 ee V-LEG sao 5 696 V-LEG 9GG 7" 696 P-LEG LEG 9% a6 a e0ul qsni UMOID ‘ . queqsIsol—="“soyy , BI10JOIA KX puog ‘* 9uBlIyy) X puog “* puog Xo BqyuRG puog x 1eyeypuey ssomQ, juDjsisal alan szuasod Yyj0q Yom 0} se0n4 Buarsyyn sayoruwa urojien pun puog Bburajoaus sassovo Us Worones ysnu Umoda sof Uornbaibos buypoos * iT L 2isvy, 164 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV The data for F, seedling segregation in crosses of Bond with Landhafer, Santa Fe, Ukraine and Victoria are presented in Table 7. Statistical tests showed good agreement between observed and expected results except in tests employing race 226 in the cross Bond x Victoria. (ii) Adult plant tests. F, generations of crosses involving Bond with Landhafer, Santa Fe and Victoria respectively were classified for mature plant field reactions and the data are presented in Table 8. With the operation of an additional partially dominant factor in Landhafer the expected F, ratio in this cross was 1005 resistant: 19 susceptible. In the cross Santa Fe x Bond a corresponding 15 TABLE 8 F, segregation for adult plant crown rust reaction in crosses involving Bond and certain varieties utilising races to which both parents were resistant Adult plant reactions Cross Total Expected ratio P value Resistant Susceptible ie R MR MS Ss) Landhafer x Bond 231 28 — 3 5 267 1005 Res.?: 19 Sus. 0-20-0-10 Santa Fex Bond 483 122 18 7 26 656 15 Res.: 1 Sus. 0-20-0-10 Bond x Victoria .. 273 15 — 2 3 293 97-4 Res.: 2-6 Sus. 0-50-0-30 1J=immune, R=resistant, MR=moderately resistant, MS=moderately susceptible, S=susceptible. 2 Res.=resistant, Sus.=susceptible. resistant : 1 susceptible ratio was expected; in the cross Bond x Victoria the expected percentage of susceptible plants was 2-6 (or seven-sixteenths of 5-92 °%). Statistical agreement between observed and expected results was good as shown in Table 8. However it was subsequently observed in progeny tests TABLE 9 F, seedling behaviour for crown rust reaction in crosses involving Bond and certain varieties utilising races to which both parents were resistant Crown Cross rust Total Expected ratio P value race Res.1Seg. Sus. Landhafer x Bond (a)? .. 29 39 = «62. 6 107 65 Res.: 152 Seg.: 19 Sus. 0-70-0-50 (O) os 2 2b9 10 «15 2 27 65 Res.: 152 Seg.: 19 Sus. 1:00-0:90 Total 49 77 8 134 65 Res.: 152 Seg.: 19 Sus. 0:70-0:50 Santa Fe x Bond (@)) a .. 259 49 62 GUN 7 Res.: 8 Seg.: 1 Sus. 0-80-0:70 (D)) ss ee 64 74 9 147 7 Res.: 8 Seg.: 1 Sus. 1-00-0-90 Total 113 136 15 264 7 Res.: 8 Seg.: 1 Sus. 0-90-0-80 Bond x Ukraine () so co Zed Ce il ale) 9 109 U leesh3 3 See, 3 Il SRE 0:50-0:30 237-4. Bond x Victoria @) sc no BACs 118} By 6 51 25-5 Res.: 62-2 Seg.: 12-3 Sus. 1-00-0-90 (GG) .. 286 27 59 16 102 25-5 Res.: 62-2 Seg.: 12-3 Sus. 0:70-0:50 Total 40 91 22 153 25-5 Res.: 62-2 Seg.: 12-3 Sus. 0:70-0:50 1 Res.=homozygous resistant, Seg.=segregating, Sus. =homozygous susceptible. 2 (a)=F, material classified for adult plant reaction; (b)=F, material classified for seedling reaction type. E. P. BAKER AND Y. M. UPADHYAYA 165 of the susceptible plants from the cross Landhafer x Bond that one segregated for the Bond type of resistance and susceptibility, one for the Landhafer type of resistance and susceptibility, whilst one was homozygous resistant for the Landhafer reaction type. This was probably due to the fact that the gene allelic to the dominant complementary factor in Bond was not operative at the adult plant stage. On this hypothesis the expected F, ratio was 249 resistant : 7 susceptible ; on the sample tested this ratio could not be statistically distin- guished from the respective 1005: 19 ratio previously hypothesized. Correlated behaviour of F, seedlings and adult plants in the cross Bond x Ukraine is presented later. (2) F, Segregation Representative samples of plants from each reaction class were tested for random F, progeny behaviour in various crosses. The observed and calculated expected frequencies of F, lines are presented in Table 9. The expected ratios in the different crosses were as follows : Homozygous Homozygous resistant Segregating susceptible Landhafer x Bond .. fe a os 85 152 19 Santa Fe or Ukraine x Boul ae A aie 7 8 1 Victoria x Bond Bs Hh a ty 25-5 62-2 12-3 Results showed good agreement between observed and expected results and confirmed assumptions that in the different crosses various factors conditioning seedling resistance were operative as follows : In the cross Bond xlLandhafer, Bond contributed two dominant complementary factors; the variety Landhafer contributed two factors one of which acted in a dominant complementary manner to one of the two factors in Bond, the second being independent and dominant. The varieties Santa Fe and Ukraine also had one independent dominant factor but possessed a complementary allele to one of the two Bond factors. The genes were allelic and not identical for reasons previously indicated. In the cross Bond x Victoria there were no interactions between the factors contributed by the two varieties and in all six factors were concerned in conditioning seedling resistant in this Cross. (d) elationship of F, or F, Segregating Reactions and Reaction Types to Different haces, Between Segregating Seedling and Adult Plants, and Between F, and I, Segregating Generations (1) F, Seedling compared with F, Seedling Inoculations were made on the primary leaf with a rust race to which both parents were resistant, and the secondary leaves inoculated with a race to which only Bond was resistant. The results from such studies involving three crosses are set out in Table 10. In all instances the susceptible classes were susceptible to both races. The majority of seedlings exhibiting an intermediate reaction type on the primary leaf showed full susceptibility on the secondary leaves, indicating that they possessed factors for resistance from the parent other than Bond. Some seedlings from this category, however, showed the Bond reaction type to races to which only Bond was resistant, indicating only partial dominance of the Bond genotype or the action of modifying genes. Considering the total resistant class the expected fractions of F, plants susceptible to the second race were thus: Cross Landhaferx Bond .. .. —57/237 Cross Santa Fex Bond .. .. —24/60. Cross Bond x Ukraine oP .. —24/60 In all three crosses the deviations were not statistically significant at the 5% probability level. STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV 166 G0-0-01-0=ONTPA dy £0E-0-OL-0=ONTBA de *0E-0-0G-0=9NTRA da ‘sgoyoerq UI sonteA poyoodx® _ 1 a oa Il = = SI == 7-§ (8°33) (Z:¥§) (3 S01) (8-91) 2(6°SF) (1 SPT) 0g LZ LOT 1ST I? OST [240.L L = = CP g jz SI = r= &% g VG cg = OFT 8G SI |) OuleIy () x puog puog < CX i eyUueg puog x doyeypue Ty ¥-8 L=8 £0 v-& i=s £0 r-8 t= etqiydeosng qZuBeysIsoy eiqiydeosng qgueqysisoy etqiydeosng JUBeYSISOYT 98G —0BYy 986 0B 986 Q0ByY (queqsisor sqguered Yy30q) soon. Burajoaur asoyy pup qunjsisas auan squauvd yzoq Yorym 07 sa0nL Buzajoauy 87891 Woamjeg sessoud PUOT Ud Sa (queqystseq puog Ajuo) sodéy uoroves * qunjsisas som puog fyuo yoryn of OL Biv, sedéy uomovoer “7 dhy uoyovas buypees *q fo uoynwossy E. P. BAKER AND Y. M. UPADHYAYA 167 One interesting observation was noted in the cross Bond x Ukraine when F, seedlings classified for reaction type to races 237-4 and 226 were reinoculated with race 286 fifty-two days after sowing. All resistant seedlings maintained their resistance whereas of the seedlings susceptible to race 226, 16 out of 28 were resistant to race 286. This shows perfect agreement with the 15-7 resistant plants expected on the assumption that the complementary factors for adult plant resistance in Ukraine were operative at this stage of growth. Presumably the third factor conditioning adult plant resistance in Ukraine was not operative against this race. (2) F, Seedling compared with F, Adult Plant Seedlings classified for reaction type in crosses of Bond with Santa Fe, Ukraine and Victoria were transplanted for observations on adult plant field reactions, and the data are presented in Table 11. The assumptions relating to gene behaviour in different crosses were : In the cross Santa Fe x Bond the factors operative in the seedling stage conditioned adult plant resistance in addition; the complementary allele in Santa Fe, operative in the adult stage only, resulted in all resistant seedlings maintaining their resistance and the susceptible seedling group proving susceptible subsequently. In the cross Bond x Ukraine, the seedling resistance Ukraine factor operated against few races only and was ineffective in conditioning adult plant resistance in the field where several races were involved. ‘The three factors in this variety conditioning only adult plant resistance were operative. The complementary allele was also effective in Ukraine (the comparable factors in Fulghum and in Santa Fe behaving similarly), the resistant seedling class producing 21 susceptible adult plants in a population of 960 and the susceptible seedling category 57 resistant plants in a population of 64. In the cross Bond x Victoria 56-25 per cent of seedlings maintained their resistance due to the operation of the Bond factors. On the 31-45 per cent resistant due to segregation of factors from Victoria, 0-55 were susceptible and of the 12-3 per cent susceptible seedlings 2-04 were susceptible as mature plants. From the data presented in Table 11 two plants in each of the crosses Santa FexBond and Bond x Ukraine did not behave exactly as expected. The reaction type class ‘‘; ’’—‘‘ 1” in the cross Bond x Ukraine was considered to possess the Ukraine type of resistance on the basis of F; progeny reaction types. No statistically significant deviations were obtained. No susceptible adult plants were observed in the Victoria type seedling reaction group ; however, since in the population examined only 0-8 was expected, this deviation was clearly not significant. (3) KF, Seedling compared with F,; Seedling Certain F, plants were classified for reaction type to races to which both parents were resistant and their F, progenies tested against similar races or against a specific race to which Bond alone was resistant. The information obtained in this regard is set out in Table 12 and shows that two plants, one each in the crosses Bond x Ukraine and Bond x Victoria, from the resistant F, category yielded susceptible progenies in tests involving races to which both parents were resistant. One line in the cross Bond x Victoria from the susceptible F, class segregated with a preponderance of susceptible plants indicating the operation of the factor Ve, and its linked inhibitor. In tests involving races to which Bond alone was resistant the only lack of correlation was that shown when a homozygous resistant line with a reaction type similar to Bond was derived from an F, plant exhibiting an intermediate reaction type. Apart from this discrepancy the behaviour of plants of an STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV 168 ‘00-T=0N[eA Jo 0L-0-08-O=enTBA gg ‘SIoquINU [[BUIS IOF poqyoor10o sonjea ,X%~< GQ-0-OL-O=onTeA gy, 0G:0-0L:0=9nTRA de ‘sjoyovlq Ul son[VA poqgoodxg ; ‘etqiuydeasns=—g ‘elqrydeosns Ajeqyerepowm—= gy ‘Jueysisor AToYeIopoUI= YP “uUeIsIsoI=y ‘ounvIWI=T _ 9(0-3) (8-0) (0:3) s(Z-&) (0-0) (0-01) (0-0) j = = G j j 6 = = I %% eg “ §-ST (0-01) (FP) (0-2) (0-91) (8-89) (0:¥#L) (0-0) (0:92) (0:19) 2(0-§L) Or GF LG SI OL GL I 9% 19 GL my “s UN ‘YW ‘WD P-§& ug — UT 1-:0 -§ ts -0 7-& AGiaAL : -0 BILOJFOLA x puodg OurIe.y () >< puog puog »*« OL B4JUBS SUOTJOBOL sedA4 uoMovor Sulfpeog qgueld 4ympy DUOPLA PUD aUIDIy ‘ag Duy ynn puog Burajoaus sassovo ue adh wouonos Il S1aviL sof paifrssnjo sbuajpoas * 7 fo suowones qunjd ynpy E. P. BAKER AND Y. M. UPADHYAYA 169 intermediate reaction type was as expected, progenies segregating for Bond genes or homozygous susceptible lines being produced, indicating the presence of factors contributed by the second parent. (4) KF, Adult Plant compared with F, Seedling The F, segregation pattern in the cross Landhafer x Bond was based on the operation of five factors, two from Bond and three from Landhafer. Of TABLE 12 F, seedling behaviour for crown rust reaction of F', plants classified for seedling reaction type in crosses involving Bond when tested against various races F, behaviour to F, reaction types to races to which races to which both races to which only both parents were parents were resistant Bond was resistant resistant Res. Seg. Sus. Res. Seg. Sus. Race 226 Race 259 Race 286 Landhafer x Bond 0; 6 10 — 1 17 5 ;-1 2, ] — 1 il 1 3-4 —- — 8 — —— 8 Race 259 Race 259 Race 286 Santa Fe x Bond 0; 26 40 = 5 58 3 ;-1 33 14 — — 3 46 2- 1 23 — — — 25 3-4 — —- 9 — — 9 Race 237-4 Races 237 and 237—4 Race 286 Bond x Ukraine 2 32 27 1 7 61 6 7 = 1) LZ 19 — — 1 47 3-4 —— — 12 — an 16 Race 237-4 Race 286 Race 259 Bond x Victoria 0; 21 36 — 9 48 2 ;-1 7 8 1 — 3 18 2— 2 16 — — — 18 3-4 — 1? 10 = — 11 1 Res. =homozygous resistant, Seg.=segregating, Sus.=homozygous susceptible. 2 Segregating with preponderance of susceptible seedlings. these one factor in Landhafer conditioned adult plant resistance only. While the susceptible F, plants were therefore expected to produce only susceptible F, seedling progenies in tests involving races to which both parents were resistant, some resistant F, adult plants when similarly tested were expected to yield susceptible seedling progenies. From data presented in Table 13, containing information pertinent to this correlated study, some F, susceptible plants gave either homozygous resistant or segregating lines due to segregation of the Bond factors effective against. race 286. These were presumed errors in F, classification or labelling. Statistically good agreement between observed and expected results in the segregation of resistant F, plants was obtained on the basis of expected ratios of 112 homozygous resistant: 608 segregating : 285 homozygous susceptible and 340 homozygous resistant: 608 segregating : 57 homozygous susceptible lines in tests involving races 286 and 259 respectively. In the cross Santa Fe x Bond two factors in Bond and two in Santa Fe, one allelic to one of the Bond factors, were effective in conditioning both seedling and adult plant resistance. Hence susceptible F, adult plants would be expected to produce homozygous susceptible F,; lines and resistant plants to yield homozygous resistant progenies or segregating lines in tests involving races to which both parents were resistant. Probably due to misclassification two in the population of 103 resistant plants produced susceptible progenies and of 170 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV the 14 susceptible plants three gave segregating progenies. The former two plants were moderately resistant and the latter three moderately susceptible and all were probably minor misclassifications. It was evident, however, that the same factors conditioned both seedling and adult plant resistance in Bond and also Santa Fe. TaBLE 13 F, seedling behaviour for crown rust reaction of F', plants from the cross Landhafer x Bond classified for adult plant reaction Adult F, behaviour to plant reactions races to which only Bond races to which both Bond was resistant and Landhafer were resistant Race 286 Race 259 Res.1 Seg. Sus. Res. Seg. Sus. ite mn hus 11 62 15 33 48 2 R-MR oe —- 2 16 2 13 3 Total Il 64 31 35 61 5 Expected 11:8 64-1 30-1 34-2 61-1 5:7 MS-S .. we — I 6 1 1 4 1 Res. =homozygous resistant, Seg.=segregating, Sus. =homozygous susceptible. 2T=immune, R=resistant, MR=moderately resistant, MS—=moderately susceptible, S=susceptible. In the cross Bond x Victoria, due to the operation of the factors from Victoria conferring mature plant resistance, only one quarter of the total population, in which the proportion of resistant plants was seven-sixteenths, was expected to give susceptible F,; seedling progenies in tests involving races to which both parents were resistant and similarly one-fifth of the susceptible F, class was expected to produce segregating F, lines. Of the 209 plants tested from the former class, 21 gave susceptible progenies compared with 21-4 expected. Since only five susceptible F, plants were progeny tested, the fact that no segregating line was observed was obviously not a significant discrepancy since only one was expected. (5) F, Seedling compared with F, Seedling In the four crosses involving Bond parentage approximately 25 seeds of each F, line were tested to one race to which both parents were resistant and a similar sample tested to a second race to which only Bond was resistant. The various frequencies and interrelationships on the genetic bases previously formulated were as follows: Behaviour to race to which Behaviour to races to only Bond was resistant Cross which both parents were resistant Homo. Homo. Res. Seg. Res. Total Landhafer x Bond .. Homo. Res. A ae 28 38 19 85 Seg. ee oo 114 38 152 Homo. Sus. ee ae — — 19 19 Total nee 28 152 76 256 Santa Fex Bond and .. Homo. Res. at 4 14 10 28 Bond x Ukraine .. A Seg. — 18 14 32 Homo. Sus. ot — — 4 4 Total 4 32 28 64. Bond x Victoria .. .. Homo. Res. ahs 6:25 10-25 9-0 25-5 Seg. = 39-75 22-5 62-25 Homo. Sus. 0 — — 12-25 12-25 Total 6-25 50-0 43-75 100-0 E. P. BAKER AND Y. M. UPADHYAYA il7ak The observed and expected frequencies, in brackets, are shown in Table 14. Since the F, population was classified for reaction type to a race to which both parents were resistant, the expected frequencies were calculated separately within the total of each reaction type class to races to which both parents were resistant and not for the overall totals. There was good agreement between observed and expected results in all crosses except Santa Fe x Bond. In every test homozygous susceptible lines to one race invariably behaved similarly to the second race. One line in each of the crosses Bond x Ukraine and Bond x Victoria segregated to the race to which both parents were resistant but was homozygous resistant to the second race. This may have been due to insufficient plants being tested to detect susceptible seedlings in a line actually segregating ; in this case the line would be designated erroneously as homozygous resistant. TaBLE 14 Correlation of F' seedling crown rust behaviour in Bond crosses between tests involving races to which both parents were resistant and those involving races to which only Bond was resistant F, reactions to F,; reactions to race to which Cross race to which only Bond was resistant Total P value both parents were resistant Res.t Seg. Sus. Race 259 Race 286 Landhafer x Bond Res. 12 (15-1)? 24 (20-6) 10 (10-3) 46 0-50-0-30 Seg. — 52 (57-8) 25 (19-2) 77 ~=0-20-0-10 Sus. — — 13 13 Santa Fe x Bond Res. 11 (13-9) 26 (48-9) 60 (34-6 97 <0-001 Seg. — 86 (75-9) 49 (59-1) 135 0-10-0-05 Sus. — . — 21 21 Race 237 and 237-4 Race 286 Bond x Ukraine Res. 6 ( 7-1) 25 (25-0) 19 (17-9) 50 0-90-0-80 Seg. 1 ( 0-0) 25 (25-3) 19 (19-7) 45 0-90-0-80 Sus. — — 16 16 Race 226 Race 259 Bond x Victoria Res. 5 ( 7-4) 17 (12-0) 8 (10-4) 30 =0-20—-0-10 Seg. 1 ( 0-0) 35 (40-2) 28 (22-8) 64 0-30-0-20 Sus. — — Il 11 1 Res. =homozygous resistant, Seg.=segregating, Sus. =homozygous susceptible. 2 Expected values in brackets. In the cross Santa Fe x Bond there was a statistically highly significant deviation from the expected results due mainly to an excess in the number of lines homozygous resistant to race 259 and homozygous susceptible to race 286 over that calculated. This is difficult to explain since the number of lines homozygous resistant to both races was close to that expected. The F; ratios in tests involving the two races independently from data presented in Tables 5 and 9 also showed good agreement on the basis of the hypotheses proposed. Association between crown rust resistance factors in Bond and stem rust resistance factors in Burke and Laggan In the crosses Burke x Bond and Laggan x Bond 228 and 235 F, lines respectively were tested against race 259 of crown rust and a mixture of races 2 and 12 of oat stem rust. The independent behaviour of genes governing crown rust resistance in Bond and the gene (Rd,) conditioning stem rust resistance in both Burke and Laggan is clear from the data presented in Table 15. B (B) 172 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV (C) Inheritance of grass clump habit and its association with the inheritance of crown rust resistance in the cross Landhafer x Bond In F, adult plants in the cross Landhafer x Bond 49 in a total of 267 plants Showed a grass clump habit of growth. These numbers agreed statistically with a ratio of 13 normal: 3 grass clump (P=0-90-0-80) suggesting the action of an inhibitor gene to a dominant gene for grass clumping. Similar gene TABLE 15 Association of F', stem rust and crown rust reactions in crosses of Bond with Burke and Laggan Stem Crown rust behaviour rust behaviour Burke x Bond Total Laggan x Bond Total Res.1 Seg. Sus. Res. Seg. Sus. Res. .. .. 6 ( 5-4)? 36 (38-2) 19 (17-4) 61 1 (1-9) 27 (25-6) 23 (23-5) 51 Seg. .. .. 10 (10:4) 70 (70-4) 38 (33:2) 118 4 (4:8) 61 (63-3) 61 (57-9) 126 SUSsaEer .. 4( 4:2) 37 (30-4) 8 (14:4) 49 4 (2:3) 30 (29-1) 24 (26-6 58 Total 20 143 65 228% 9 118 108 2354 1 Res.=homozygous resistant, Seg.=segregating, Sus.=homozygous susceptible. 2 Hxpected values in brackets based on marginal totals. 3 y7=3-56; P=0-50-0-30. 4 yi=2-43; P=0-70-0-50. action for grass clump habit has been observed in wheat crosses (McMillan, 1937 ; Watson, 1943). The independence of genes determining this character from those conditioning adult plant resistance in Bond and Landhafer is shown from the data in Table 16. A number of weak F, plants were observed and TABLE 16 Association of adult plant reaction to crown rust and grass clump habit in the F, generation of a cross between Bond and Landhafer Adult plant Growth habit reactions 1 Grass clump Total Normal Weak ie a6 Be < 147 (147-1)? 40 (41-5) 44 (42-4) 231 R-MR a oun 17 (17-8) 7 (5-0) 4 (5-2) 28 MS-S .. a0 = 6 (5-1) 1 (1-5) 1 (1¢4) 8 Total 218 49 2678 Expected ratio 216-9 50-14 (13 normal : 3 grass clump) 17=immune, R=resistant, MR=moderately resistant, MS=moderately susceptible, S=susceptible. 2 Expected values in brackets based on marginal totals. 2 x3 for independence=1-67; P=0-90-0-80. 4 yi=0-03; P=0-90-0-80. these may have been heterozygous for the inhibitor gene in the presence of the dominant factor for grass clumping, indicating partial epistasis. DISCUSSION AND CONCLUSIONS From the present studies it is evident that the immunity of Bond to Australian races of crown rust in both seedling and adult plant stages was conditioned by two dominant complementary genes. However, dominance was incomplete at high temperatures and the genotype heterozygous for both factors E. P. BAKER AND Y. M. UPADHYAYA 173 gave a moderately susceptible reaction type modifying the characteristic 9 resistant: 7 susceptible F, ratio to a respective 5: 11 ratio. These findings are in accordance with the observations of other investigators, including Hayes et al. (1939) and Weetman (1942). It is proposed to designate these factors as Bd, and Bd»p, the line below each symbol indicating complementary gene action. Presutnably one gene was contributed by each of the parents, Golden Rain and Red Algerian selection, to the hybrid from which it was selected. Segregation in a cross between Bond and Golden Rain is currently being studied. to investigate this hypothesis. Operation of a single factor was observed in a number of intervarietal crosses involving Bond. In its crosses with Algerian and Fulghum at the mature plant stage monogenic segregation was observed, whilst in crosses with Santa Fe and Ukraine, involving races to which these varieties were also resistant, duplicate factor segregation was noted. Hayes et al. (1939) assumed the presence of a common factor in such crosses, Finkner et al. (1955) only a Single factor in both varieties whilst Osler and Hayes (1953) and Weetman (1942) detected the presence of factors in Santa Fe and Ukraine allelic to one of the Bond genes. The present studies confirmed the latter findings since the complementary factors in Bond were operative in the seedling stage against races to which the second parent was susceptible. It was also evident that the alleles in the varieties Algerian, Fulghum, Santa Fe and Ukraine were not identical since they were operative only against the races to which the respective parents were resistant, either in both seedling and adult plant stages or in the mature plant stage only. These studies extended the genes for crown rust resistance identified in certain varieties. The complete genotypes of these varieties for crown rust resistance are given below, where non-interacting genes are indicated by numerical suffixes, interacting (complementary) genes given alphabetical suffixes and allelomorphic genes indicated by apostrophe (’) marks according to increasing order of reaction type or reaction. Torrie (1939) claimed that Bond possessed one strong gene and that another masked its effect. If Torrie’s strong gene in Bond is that designated as Bd, the alleles in the present instance would be relative to the second Bond factor Bdp. Those in Santa Fe, Ukraine and. Fulghum would in the proposed terminology be designated as Bdp’, Bdy”’ and Bdp’” respectively. Similarly the alternative gene in Landhafer, assumed to act in complementary fashion with Bd, would be designated Bdy. This factor was also operative against race 286 to which Landhafer was ‘susceptible. The complete list of genes, including those documented by Upadhyaya and Baker (1965), in varieties in which additional factors were revealed in Bond crosses are thus: Landhafer—three factors. One factor (Lid,) conditioned seedling as well as adult plant resistance. A second factor Ld, was responsible for adult plant resistance only and the third factor Bdy, was an alternate complementary factor: to one (Bd,) of the two complementary Bond factors in conditioning seedling resistance only. Santa Fe—three factors. One factor (Sf,) conditioned seedling as well as adult plant resistance. A second factor (Tr.) acting in complementary fashion with Trp in Trispernia inhibited the action of Sf, against race 203 in the seedling stage only. The third factor Bdy’ was complementary in action with Bd, and allelic to Bdp. This factor conditioned seedling resistance to races to which Santa Fe was resistant and also resistance in the adult plant stage. Ukraine—five factors. One factor Sf,’, allelic with Sf,, conditioned only seedling resistance to race 237. Three genes Mu,, Mua and Mup, conferred. 174 STUDIES ON THE INHERITANCE OF RUST RESISTANCE IN OATS, IV adult plant resistance only, the two latter factors acting in complementary fashion. The fifth factor Bd,” was a member of the allelic series at the Bdp locus. Fulghum and | Jeenterm_aa gene. The Bdy’” member of the allelic series at this locus acted in complementary fashion with the gene Bd, to OSE adult plant resistance only. The genes previously identified in Trispernia were Sf,”, allelic with Sé,, but exhibiting a higher reaction type, and Trp complementary to Tra. No variety used in these studies or those previously reported in this series of papers was resistant to all Australian races of crown rust and the allelism indicated is obviously a barrier in attempts to combine genes from different varieties to provide a more comprehensive genetic basis for resistance in breeding commercial varieties. The order of epistasis or dominance in the varieties studied in this series of papers was Bond (immune) ; Landhafer, Santa Fe, Ukraine (highly resistant) ; Trispernia, Victoria (resistant) ; Fulghum, Algerian, Burke, Laggan (susceptible). Plants of a grass clump habit were observed in the F, generation of the cross Bond x Landhafer. Coffman (1964) detected dwarf segregates, some apparently of grass clump or grass tuft habit, in certain intervarietal crosses. He con- sidered dwarfing in general to be due to one, two or three gene pairs with various types of interaction when more than one gene was implicated. References Baker, H. P., and Upapuyaya, Y. M., 1955.—Physiologic specialisation of crown rust of oats. Proc. Linn. Soc. N.S.W., 80: 340-357. Cocuran, G. W., JoHNSON, C. O., HyENE, E. G., and Hansine, E. G., 1945.—Inheritance of reaction to smut, stem rust and crown rust in four oat crosses. Jour. agric. Res., 70: 43-61. Corrman, F. A., 1964.—Inheritance of morphologic characters in Avena. United States Dept. Agric. tech. Bull. No. 1308. FINKNER, V. C., 1954.—Genetic factors governing resistance and susceptibility of oats to Puccinia coronata Corda variety avenae, F. & L., Race 57. Lowa State Coll. agric. Expt. Stat. Res. Bull., 411: 1040-1063. Grirritus, D. J., 1953.—Varietal resistance and susceptibility of oats to crown rust. Plant Pathology, 2: 73-77. Hayes, H. K., Moors, M. B., and Staxman, E. C., 1939.—Studies of inheritance in crosses between Bond, A. byzantina and varieties of A. sativa. Minn. agri. expt. stat. tech. Bull. 137. Kear, W. R., and Hays, H. K., 1950.—Studies of inheritance in crosses between Landhafer, A. byzantina L. and two selections of A. satwa L. Agron. Jour., 42: 71-78. Ko, S. Y., Torriz, J. H., and Dickson, J. C., 1946.—Inheritance of reaction to crown rust and stem rust and other characters in crosses between Bond, A. byzantina, and varieties of A. sativa. Phytopath., 36: 226-235. LitzENBERGER, S. C., 1949.—Inheritance of resistance to specific races of crown rust and stem rust, to Helminthosporium blight and of certain other agronomic characters of oats. Jowa agri. Expt. Stat. Res. Bull. 370. McMinnan, J. R. A., 1937.—Investigations on the occurrence and inheritance of the grass clump character in crosses between varieties of J. vulgare (Vill.). Coune. sci. industr. Res. Melbourne, Bull. 104. OstER, R. C., and Hays, H. K., 1953.—Inheritance studies in oats with particular reference to Santa Fe type of crown rust resistance. Agron. Jour., 45: 49-53. ‘Torriz, J. H., 1939.—Correlated inheritance in oats of reaction to smuts, crown rust, stem rust and other characters. Jour. agric. Res., 59: 783-804. Urapuyaya, Y. M., and Baxerr, H. P., 1960.—Studies on the mode of inheritance of Hajira type stem rust resistance and Victoria type crown rust resistance as exhibited in crosses involving the oat variety Garry. Proc. Linn. Soc. N.S.W., 85: 157-179. , , 1962a.—Studies on the inheritance of rust resistance in oats. I. Inheritance of stem rust resistance in crosses involving the varieties Burke, Laggan, White Tartar and Anthony. Proc. Linn. Soc. N.S.W., 87: 141-147. , 1962b6.—Studies on the inheritance of rust resistance in oats. II. The mode of inheritance of crown rust resistance in the varieties Landhafer, Santa Fe, Mutica Ukraine, Trispernia and Victoria in their crosses with susceptible varieties. Proc. Linn. Soc. N.S.W., 87: 200-219. E. P. BAKER AND Y. M. UPADHYAYA 175 Urpapuyaya, Y. M., and Baxer, E. P., 1965.—Studies on the inheritance of rust resistance in oats. III. Genetic diversity in the varieties Landhafer, Santa Fe, Mutica Ukraine, Trispernia and Victoria for crown rust resistance. Proc. Linn. Soc. N.S.W., 90: 129-151. WateERHOUSE, W. L., 1938.—Some aspects of problems in breeding for rust resistance in cereals. Jour. roy. Soc. N.S.W., 72: 7-54. , 1952.—Australian Rust Studies. IX. Physiologic race determinations and surveys of cereal rusts. Proc. Linn. Soc. N.S.W., 77: 209-258. Watson, I. A., 1943.—Inheritance studies with Kenya varieties of Triticum vulgare Vill. Proc. Linn. Soc. N.S.W., 68: 72-90. Wertman, I. M., 1942.—Genetic studies in oats of resistance to two physiologic races of crown rust. Phytopath., 32: 19 (abst.). A NEW CORALLANID ISOPOD PARASITIC ON AUSTRALIAN FRESHWATER PRAWNS HK. F. RIEK Division of Entomology, C.S.I.R.O., Canberra, A.C.T., Australia [Read 28th September, 1966] Synopsis A second species of Austroargathona is described from inland New South Wales and Queensland. The species is parasitic on freshwater prawns of the genera Macrobrachium (Palaemonidae) and Paratya (Atyidae). The type species and only other described species is parasitic on the related prawns of the easterly flowing streams of coastal Queensland and New South Wales. AUSTROARGATHONA PICTA, Sp. NOV. (Figs 1-6) Non-ovigerous Female. Length up to 11 mm. Very similar in general appearance to caridophaga Riek (Riek, 1953) but differing a little in colour intensity and pattern (Fig. 1). Median chromatophore band of peraeon divided {p Yo a Prom Orr Wy Fig. 1. Awstroargathona picta sp. nov., Dorsal view x 9. except on the fourth segment so that there are six indistinct bands except on this one segment. The median two bands of the telson have almost merged so that the telson appears to have a single very wide median band, a distinct, PROCEEDINGS OF THE LINNEAN SocrETy oF New South Watss, Vol. 91, Part 3 BE. F. RIBK AA wide, light laterad area and narrow dark margin. The ramose chromatophores are uniformly pigmented and have sharply defined margins. The cephalon is relatively wider and the eyes less prominent and the lateral margins of the telson are more uniformly convex than in caridophaga. Mouthparts distinctive, mandible and second maxilla differing most from those of caridophaga. First and second antenna very similar to those of caridophaga, but first article of peduncle of first antenna rather more expanded at base anteriorly ; frontal lamina wedge-shaped with a slightly concaved anterior margin and tapering to a rounded point posteriorly ; mandibular palp (Fig. 2) with basal Figs 2-6. Austroargathona picta sp. nov. 2. Mandible. 3. First maxilla. 4. Second maxilla. 5. Maxilliped. 6. First peripod. segment expanded, wider than the following segment, second segment with two Series of apical spines, an outer marginal series of seven stout subequal spines, the second the longest, the others gradually shortening distally, but each at least as long as the width of the segment, a very small spine proximal to this Series, also a series of three apical spines on the ventral side of the segment, the anterior one much the longest and as long as those of the marginal series, posterior margin of apical segment with a series of about eighteen setae, the distal three very long and stout, with the middle one of these three almost as long as the segment; mandible without a well developed cutting edge, apex 178 A NEW CORALLANID ISOPOD produced into an acute, stout, dark-coloured spine and bearing a series of four fine spines on the median side at the base of the terminal spine, the innermost spine the finest. These spines are less than half the length of the terminal spine and are very fine and transparent. In the description of caridophaga the mandible and its palp were inadequately described. They are similar to those in the present species, but the apical spine of the mandible is relatively smaller, and there are apparently only two shorter but stouter mesal spines on the much more narrowed apical lobe. First maxilla (Fig. 3) with tapering base, outer lobe tapering to the acute, distinctly curved, terminal unguis, inner lobe small, rather narrow and with incurved, apex; second maxilla (Fig. 4) small, anterior margin rounded, slightly expanded on the median side (in caridophaga the muscle fibres of attachment were confused with the basal structure of the second maxilla so that the structure in the two species is more similar than would appear from the original figures). Coxal plates not quite as acutely pointed and their outer margins more crenulated than in caridophaga. Maxilliped (Fig. 5) and first periopod (Fig. 6) similar to those of caridophaga but palp of maxilliped more clearly 4-segmented. Types.—Holotype non-ovigerous female and paratype non-ovigerous females in the Australian Museum Collection. Type Locality.—Darling River at Bourke, N.S.W. Distribution.—Darling River at Bourke, N.S.W. (7 Dec. 1959, V. McCristal, 29 Mar. 1960, N. E. Milward and 17 Mar. 1961, J. A. Bishop); Darling River near Gundabooka Bridge (18 Mar. 1961, J. A. Bishop); Darling River, 140 miles downstream from Bourke (18 Mar. 1961, J. A. Bishop); Darling River near Tilpa (19 Mar. 1961, J. A. Bishop) ; Bogan River, 2 miles c. west of Tarcoon, N.S.W. (23 Mar. 1960, N. E. Milward) ; Condamine, Q’ld. (10 Apr. 1957, EH. F. Riek); Namoi River, near Narrabri, N.S.W. (May, 1961, W. D. Williams) ; Boonoo Boonoo River, near Tenterfield, N.S.W. (24 May 1961, W. D. Williams), parasitic on cat-fish. Hosts.—Kctoparasitic on the freshwater prawns Macrobrachium australiense cristatum Riek (Palaemonidae) and an undescribed species of Paratya (Atyidae). Only at Condamine did the species occur on Paratya. The host prawns are restricted to freshwaters. The cat-fish is very probably not a normal host of the species. The paratypes are all considerably smaller than the holotype. They differ in having only a single fine spine on the median side of the mandible. It is considered that this species of Austroargathona will be found to be widespread in the inland flowing streams of Queensland and New South Wales, for during flooding there is extensive interconnection between the streams. At least one of the host prawns is widespread. Reference Riek, E. F., 1953.—A corallanid isopod parasitic on freshwater prawns in Queensland. PrRoo. Linn. Soc. N.S.W., 77: 259-261. NEW AUSTRALIAN CAVE CARABIDAE (COLEOPTERA) B. P. Moore C.S.1.R.0O., Canberra [Read 28th September, 1966] Synopsis Three new species of cave-dwelling Carabidae are described, viz. Idacarabus cordicollis (Tasmania), Thenarotes speluncarius (Nullarbor Plain) and Anomotarus subterraneus (South Queensland). A. subéerraneus is of interest in providing a link between the cavernicolous Speotarus species and related, surface-dwelling lebiines. Fig. 1. Idacarabus cordicollis sp. n., paratype male. PROCEEDINGS OF THE LINNEAN SocreTy of New SoutH Watss, Vol. 91, Part 3 180 NEW AUSTRALIAN CAVE CARABIDAE (COLEOPTERA) Subfamily MERIZODINAE IDACARABUS CORDICOLLIS, sp. 0. (Figs 1-2) Elongate-oval; subapterous; dark piceous, the appendages lighter. Head ovoid, rather small; eyes small, not prominent ; labrum trapezoidal, sexsetose ; mandibles slender, acutely pointed ; antennae rather long, pubescent from the middle of the third segment. Pronotum cordiform, slightly transverse ; anterior and posterior marginal setae present, the latter at the hindangles. Elytra oval, lightly striate; no scutellary strioles; humeri effaced; third intervals with two setigerous pores, situated near the third striae. Legs rather long, slender ; male anterior tarsi with two basal segments expanded and inwardly O-2 mm. Figs 2-3. Aedeagi in left lateral view. 2. Idacarabus cordicollis sp. n. (parameres detached). dentate. Aedeagus (Fig. 2) with a slender median lobe ; parameres dissimilar, the left triangular, trisetose apically, the right styloid, quinquesetose apically. Length, 5-2-6-4 mm.; max. width, 2-:3-2-6 mm. Type (male) and allotype (female).—Tasmania, Newdegate Cave, near Hastings, 13.x1.63 (EH. Hamilton-Smith), in the South Australian Museum. Four paratypes (both sexes, one immature) : King George V Cave, near Hastings, 14.xi.63 (E. Hamilton-Smith) in the South Australian Museum and in the author’s collection. This species is, in reality, the second known member of the genus, for flavipes Lea, referred originally and tentatively by its author to Idacarabus, has now proved to be a trechine and must therefore be removed to another genus. Idacarabus cordicollis is less obviously cave-adapted than I. troglodytes Lea, the generic type species from Ida Bay Caves, and the two may readily be separated as follows :— Form elongate ; pronotum elongate, lozenge-shaped, the posterial marginal seta wanting ; eyes very small; colour castaneous ; length 6-5—7-5 mm. .. troglodytes Lea Form less elongate; pronotum slightly transverse, cordate, the posterior marginal seta present at hindangle; eyes larger; colour piceous; length Oe DEO A MAMA aces es se Goes aos eu 6 sisal Dialer ere aan Tote oe Reem RSS Mscee ones cordicollis sp. n. B. P. MOORE 181 Subfamily HARPALINAE THENAROTES SPELUNCARIUS, sp. 0D. (Figs 3-4) Mostly brownish-testaceous, the appendages lighter; eyes dark; wings well developed. Head rather small; eyes prominent, lightly inclosed behind; labrum rectangular, sexsetose ; mandibles short, acutely pointed; antennae slender, of average length for the subfamily. Pronotum slightly transverse; anterior angles well marked, subprominent; posterior angles obtuse; sides evenly rounded on front third, then lightly and obliquely contracted to base; base and apex subtruncate ; basal foveae scarcely apparent ; anterior marginal seta present at front quarter ; posterior seta wanting. Elytra elongate-oval, widest a little behind middle, regularly but lightly striate; striae unpunctured ; no scutellary strioles; third intervals with one setiferous pore, situated against [VE A ee ND O°5 mm. Fig. 3. Thenarotes speluncarius sp. n. the second stria. Legs rather long, slender; tarsi setose above ; male anterior and intermediate tarsi expanded and spongiose beneath. Aedeagus (Fig. 4) stout ; median lobe tubular, with a dorsal orifice, the apex unarmed ; parameres conchoid. Length, 5-6 mm.; max. width, 2-2-3 mm. Type (male).—Western Australia, Abrakurrie Cave (Nullarbor Plain), 13.1.64 (P. Aitken), in the South Australian Museum. Fifteen paratypes (both sexes), same locality as type, various dates (1960-64) (P. Aitken and E. Hamilton- Smith) in the South Australian Museum and the author’s collection. The placing of this new cavernicolous species in Thenarotes Bates (type Species tasmanicus Bates) must be regarded as provisional, pending an overall revision of the Australian Harpalinae. However, 7. speluncarius certainly falls within the tribe Pelmatellini, to which the cavernicolous Notospeophonus species also belong, but it differs from the latter in possessing setose tarsi and an aedeagus of entirely different form, with no spatulate apex to the median lobe (c.f. Moore, 1964, for aedeagal characters of Notospeophonus). Thenarotes speluncarius has also been collected in Buckalowie Cave and Cave Number 11, of the South Australian portion of the Nullarbor Plain ; seven surface-dwelling species, referable to the genus, range over most of the eastern States. 182 NEW AUSTRALIAN CAVE CARABIDAE (COLEOPTERA) \ AN \ KE AG Fig. 4. Thenarotes speluncarius sp. n., paratype female. B. P. MOORE 183 Subfamily LEBONAE ANOMOTARUS SUBTERRANEUS, sp. 0. (Fig. 5) Form short, depressed, subparallel; fully winged; head, pronotum and appendages brownish-testaceous ; eyes black ; elytra mostly piceous but humeri widely and margins narrowly brownish-testaceous. 5 Fig. 5. Anomotarus subterraneus sp. n., paratype female. Head large, broad, depressed; eyes large and prominent, inclosed in swollen orbits ; neck pronounced ; labrum subrectangular, transverse, sexsetose ; mandibles short, broad, strongly curved and acutely pointed ; antennae short, pubescent from the third segment. Pronotum very transverse, widest at front third ; sides strongly rounded from anterior angles to widest part, then strongly contracted and slightly sinuate to posterior angles; anterior and posterior 184 NEW AUSTRALIAN CAVE CARABIDAE (COLEOPTERA) marginal setae present; anterior angles scarcely apparent; posterior angles obtuse but well marked. Elytra rather short, subparallel, striate, the striae unpunctured ; scutellary strioles wanting; humeri rounded; third intervals with two setiferous pores, one submedian, the other subapical ; apex of abdomen bisetose in male, quadrisetose in female. Legs short but slender ; male anterior tarsi lightly dilatate and squamose beneath ; claws pectinate. Aedeagus of the normal lebiine type, scarcely differing from those of species of Speotarus (Moore, 1964). Length, 4-6-5 mm.; max. width, 1-8-1-9 mm. Type (male).—South Queensland, Riverton Cave (in bat guano), 8.11.64 (HE. Hamilton-Smith), in the South Australian Museum. Nine paratypes (both sexes, same data as for type) in the South Australian Museum and in the author’s collection. This pretty little species may be recognized within its genus by virtue of its short broad form, large head, and characteristic elytral colour pattern. It is the first-discovered cavernicolous member of the genus and, as such, is of special interest in providing an evolutionary link between the surface-dwelling lebiines and the (so far as is known) exclusively cavernicolous species of Speotarus. However, although differing markedly in build from most of its congeners, A. subterraneus shows no obvious adaptation to the cave environment. Indeed, the prominent eyes and fully developed wings suggest that the beetle may not be an obligate cavernicole. Acknowledgement The author is indebted to Mr. EH. Hamilton-Smith for the opportunity to study the interesting material dealt with in this paper. References Lea, A. M., 1910.—On some Tasmanian cave-inhabiting Beetles. Yasm. Nat., 2: 53-58. Moors, B. P., 1964.—New cavernicolous Carabidae (Coleoptera) from mainland Australia. J. ent. Soc. Q’ld., 3: 69-74. ATOPOZOA DEERATA (SLUITER): A DISCUSSION OF THE RELATIONSHIPS OF THE GENUS AND SPECIES PATRICIA KoTT (Mrs. W. B. MATHER) University of Queensland, Australia [Read 26th October, 1966] INTRODUCTION The genus Atopozoa Brewin 1956 was established for the reception of Atopozoa marshit Brewin 1956. The present discussion of the genus and its relations with other genera of the family CLAVELINIDAE Forbes and Hanly is based on a study of larval form, adult colony and zooids of Atopozoa deerata (Sluiter), taken in the Gulf of Carpentaria, North Australia, by the C.S.I.R.O. Division of Fisheries and Oceanography while conducting a Prawn Survey in the area. DESCRIPTION ATOPOZOA DEERATA (Sluiter) Distoma deerata Sluiter, 1895, p. 167 ; Polycitor coalitus Sluiter, 1909, p. 23 ; Sigillina (Polycitor) coalita, Michaelsen, 1930, p. 484; Szgillina deerata, Hastings, 1931, p. 87. Records.—Thursday I., Torres Strait (Sluiter, 1895), Malaysia (Sluiter, 1909), Great Barrier Reef (Hastings).. Material Examined.—Gulf of Carpentaria (Prawn Survey St. 1121, C.S.I.R.O. portion only of one colony). Colony.—Soft, gelatinous ; maximum circumference 6 cm., height 8 em. Surface of colony with furrows up to 1-0 cm. deep dividing surface into irregular areas, or shallow lobes about 4 cm. diameter. Two common cloacal openings observed near apex of colony in furrows protected by overhang of surface lobes. Zooids accumulated in oval-circular areas especially in surface of lobes but also present in base of clefts between lobes. Common cloacal canals posterior to zooids opening into extensive cloacal Spaces separating outer zooid bearing layer of test from central very soft test core. Zooids may be arranged in lines along either side of cloacal canals although this is rather obscure particularly from the surface due to excessive crowding and overlapping of zooids. Zooids.—(Text-fig. 1). Small, 3 mm. long, excluding long terminal branchial siphon which is as long as thorax. Atrial siphon 6-lobed, long, from posterior third of thorax, directed posteriorly. Zooids arranged in test at angle to surface, with endostyle uppermost, atrial siphon opening into cloacal canals. Three rows of 10 stigmata, anterior row with progressively shorter stigmata dorsally, Stomach small, spherical, half way down abdomen. A long vascular stolon present, with mesodermal septum. However, no muscles present here as suggested by Michaelsen (1930). Developing embryos at different levels up oviduct, eventually developing to maturity in oviducal pouch, stalked, posterior to atrial siphon ; small diverticulum of oviduct in pouch of body wall in mature zooid for reception of embryo as in Distaplia bermudiensis (Berrill, 1948, }, Text-fig. 5). PROCEEDINGS OF THE LINNEAN Society or NEw SoutH Watzs, Vol. 91, Part 3 186 ATOPOZOA DEERATA (SLUITER) Larva.—(Text-figs 2-4). Almost circular, 0-8 cm. long. Papillae carried in vertical line on frontal plate, connected to rest of body of larva by ventral stalk. Papillae unusual, consisting of central circle of adhesive cells depressed into a concavity in the frontal plate which concavity forms a cup like sucker around the adhesive cells. The adhesive cells themselves in mature larvae are carried forward on a stalk but the epidermal cup remains as a concavity in the frontal plate and is never stalked. 1.—Zooid. i Din SO VOd00 00 v ae 0 OnLy): 2.—Frontal plate showing papillae developing. 3.—Lateral view of frontal plate showing papillae. 4.—Mature larva, papillae extruded. DISCUSSION The specimen differs from those of Hastings and Sluiter only in the number of common cloacal openings. The colony and zooids also resemble those of Sigillina vasta Millar 1962 (also Millar, 1963). PATRICIA KOTT 187 Although the colonies differ and the zooids open directly to the surface, lacking a long atrial siphon and common cloacal system, the zooids of the present species resemble those of : Atopozoa marshit Brewin, 1956; Hudistoma vitreus (Sars) 1851 (see also Berrill, 1948, Millar, 1963b); Hudistoma mobiusi (Hartmeyer) 1905 (see also Hartmeyer, 1912, Millar, 1962); Hudistoma fantasiana Kott, 1957. There has been some question on the number of rows of stigmata in Hudistoma vitreus, however Millar (1963b) has confirmed the presence of three rows of stigmata in the branchial sac. Berrill (1948) had indicated four rows of stigmata in the larval form, however from his figure it appears that the most anterior row may be from the other side of the branchial sac. All the other species mentioned above have three rows of stigmata. These species, all with three rows of stigmata are unique in the Polycitorinae by virtue of a brood pouch. Larvae are known for all species except EH. mobiusi and are all similar : the papillae are not stalked, or only barely stalked, the adhesive cells being accommodated in concavities in the frontal plate as described above for EH. deerata. In the larvae of H. vitreus and EH. deerata the adhesive cells are maintained in a circular area but in 8S. vasta, HL. fantasiana and in A. marshii the adhesive area has been vertically elongated in a corresponding elongation of the ectodermal concavity and in the latter two species the number of papillae is reduced to two. In larvae of H. vitreus and A. marshiv the frontal plate has not been described as stalked, however it is probable that in more mature larvae it does become so as in Distaplia spp. (see Berrill, 1948) for Distaplia bermudiensis, Text-figs 11, 12). In mature larvae of S. vasta, and HL. fantasiana the papillae are always carried forward on a stalked frontal plate. The larvae of this group of species are therefore unique in the Polycitorinae due to the stalked frontal plate and sessile papillae. There are variations in the colonies: S. vasta and H. deerata have colonial systems; H. vitreus, A. marshi and some specimens of HL. mobiusi (Hartmeyer, 1912) have similar colonies with the zooids arranged around a stalked head ; E. fantasiana and other colonies of HL. mobiusi have unstalked investing or cushion- like colonies. Despite these variations the close relationships of zooids and larvae indicate that these species belong in the same genus and the definition of the genus Atopozoa Brewin, has been amended to include this group of six closely related species, distinct from other Polycitorinae. The genus has strong affinities with Holozoinid genera where the larval papillae are also carried forward on a stalked frontal plate, where the papillary stalk is short and thick, and where the larvae develop in a brood pouch. The present group of species differs from the Holozoinae in that they do not always form systems, zooids in many of the species opening to the exterior instead of into a common cloacal system: larval papillae are present in a vertical row and have lost a primitive triradiate arrangement; they have three instead of four rows of stigmata; the atrial siphon persists and has not been modified into a languet. Nevertheless these species demonstrate stronger affinities with the Holozoinae than with the Polycitorinae where they have been accommodated, due only to the presence of three rows of stigmata and an atrial siphon. Budding in the Holozoinae is from an epicardial extension into the vascular process (Berrill, 1948) from the posterior end of the abdomen. This enlarges when vegetative reproduction occurs. Hastings (1931) suggests this as the mechanism of budding in the present species which would be expected if the genus does belong with the Holozoinae ; rather than budding by the primitive abdominal strobilation of the Polycitorinae. This would explain the variations in the magnitude of the vascular appendage (Sluiter, 1909, p. 25) and suggests Cc 188 ATOPOZOA DEERATA (SLUITER) the possibility that certain species of other genera of Polycitorinae, e.g. Sigillina and Hyperiodistoma, characterized by a well developed vascular stolon, may also belong in the Holozoinae. It is considered that the genus Atopozoa as defined below, and as discussed above is closely related to species of the Holozoinae, and well accommodated in that subfamily. The definition of the subfamily Holozoinae, has been amended (below) to include the genus Atopozoa. CLASSIFICATION The definitions of the subfamily Holozoinae Berrill and genus Atopozoa Brewin are modified to accommodate the species as discussed above as follows : Subfamily HoLozornak Berrill, 1950 Budding from epicardial extension into posterior abdominal vascular extension ; larvae develop in stalked brood pouch; larval papillae from stalked frontal plate ; zooids with or without atrial siphon; common cloacal system present or zooids open directly to surface. Genus ATOPOZOA Brewin, 1956 Atrial siphons present ; zooids open independently or into common cloacal system ; three rows of branchial stigmata; larval papillae sessile. Type Species—Atopozoa marshu Brewin, 1956 Unfortunately, in the absence of larvae and brood pouch there are no characters of the adult or colony (unless systems are formed) which can distinguish the species from Polycitorinae unless it can be shown that an enlarged vascular stolon indicates that budding always occurs from this area, and this can be correlated with the presence of typical larvae developing in a true brood pouch from the postero-dorsal corner of the thorax. In mature zooids a rudimentary brood pouch is present before it is occupied by the developing embryo. References BeERRILL, N. J., 1948a.—Tadpole larvae of ascidians Polycitor, Huherdmania, Polysyncraton. J. Morph., 82: 355-364. , 1948b.— Budding and the reproduction cycle of Distaplia. @Q. JI microsc. Sci., 89: 253-289. , 1950.—The Tunicata. Ray Soc. Publs., 133: 1-354. Brewin, BEeryt, 1956.—Atopozoa marshi, a compound Ascidian from Western Australia. J. Proc. r. Soc. West. Aust., 40 (1): 31-32. HarrMever, R., 1905.—Ascidians from Mauritius. Zool. Jb., Suppl. viii: 383-406. —, 1912.—Die Ascidien der Deutschen Tiefsee Expedition. Wiss. Hrgebn. dt. Tiefsee- Exped. ‘ Valdivia’, 16 (H.3): 223-392. Hastines, A. B., 1931.—Tunicata. Scient. Rep. Great Barrier Reef Hxped., 4 (3): 69-109. Kort, P., 1957.—Ascidians of Australia 11. Aplousobranchiata Lahille; Clavelinidae Forbes and Hanly and Polyclinidae Verrill. Aust. J. mar. freshwat. Res., 8 (1): 64-110. MicHAELsEN, W., 1930.—Ascidiae Krikobranchiae. Fauna stidwest-Aust., 5 (7): 463-558. Mittar, R. H., 1962.—Further descriptions of South African Ascidians. Ann. S. Afr. Mus., XLVI (VII): 113-221. , 1963a.—The larva and affinities of the ascidian Sigillina vasta Millar. Ann. Mag. nat. Hist., VL :‘(18): 204-207. , 1963b.—The structure and systematic position of the Ascidian Distomum vitreum Sars. Ann. Mag. nat. Hist., VI (13): 386-388. Sars, M. (1851).—Beretning om en i Sommeren 1849, foretagen zoologiske Reise 1 Lofoten og Finmarken. Nyt Mag. Naturvid., 6: 121-211. Sturtrer, C. P., 1895.—Tunicaten in Semon, R. Zoologische Forschingsreisen in Australien und den malagischen Archipel. Denkschr. med.—naturw. Ges. Jena., 8: 163-166. , 1909.—Die Tunicaten der Siboga Expedition Pt. 2. Die merosomen Ascidien. Siboga EHxped., 56b: 1-112. INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS HK. P. BAKER Department of Agricultural Botany, The University of Sydney [Read 26th October, 1966] Synopsis The inheritance of resistance to race T—1 of bunt (Tvlletia caries (DC.) Tul. and 7. foetida (Wallr.) Liro) was investigated in crosses involving two resistant selections, Turkey C.I. 10095 and C.I. 10097, which were originally made from Turkey Red or South Dakota No. 144 wheat. Studies of F, and F,; generations were made for per cent of bunted plants in crosses between each of these varieties with the susceptible variety Baart and with the resistant varieties Martin, Turkey 3055, Rio and Selection 1403 which are testers for the M, T, R and H major genes respectively, and with the resistant variety Turkey C.I. 10015, possessing only the weak genes X and Y. Both Turkey C.I. 10095 and C.I. 10097 were found to possess weak genes for resistance. To invoke a minimum number of genes to satisfy analyses in all crosses a minor revision of the genotype for Turkey C.I. 10015, as published previously by El Khishen and Briggs, was necessary in that the gene X, and not Y, was considered linked to the HMRT linkage group- On this revised genotype for Turkey C.I. 10015, selection C.I. 10095 was considered to have the major gene R and the two minor genes X and Y, hence the genotype RRXX YY with regard to bunt resistance, whilst the resistance of C.I. 10097 was considered due to the additive or complementary effect of two weak genes, X, and an independent previously unidentified factor, W, which permitted 37-5% of bunted plants when homozygous and 47-5% when heterozygous. The most satisfactory crossover value for the R and X genes considering all crosses was approxi- mately 10% and the order of linked genes conditioning bunt resistance was determined from appropriate tester crosses as HMXRT. INTRODUCTION Bunt of wheat (Tilletia caries (DC.) Tul. and T. foetida (Wallr.) Liro) can generally be effectively controlled by seed treatment. However, Kuiper (1965) described a race of bunt which was not controlled by hexachlorobenzene, the active constituent of several seed-dressing products, and generally accepted throughout the world as one of the most effective fungicides for the control of the disease. This emphasizes the importance of the second method for the control of the disease which is by breeding resistant varieties. The bunt. organisms exhibit physiological specialization and no single gene has been identified which will control all races. However Briggs and Holton (1950) showed that a combination of two genes will give effective protection against at least 25 races. In breeding resistant varieties a knowledge of the mode of inheritance of resistance shown by different genes is essential. Investigations on the genetics of resistance to a specific race of bunt will enable resistance genes to be documented and their mode of inheritance studied. As new bunt races are discovered the behaviour of varieties of known genotype can then be tested. Different genes may be required in breeding to incorporate resistance against groups of races of varying pathogenicity to afford protection against all prevalent races. Briggs and his associates at the College of Agriculture, Davis, California have studied the inheritance of resistance in 18 wheat varieties to race T-1 of 7. caries and in the course of these investigations at least seven genes for resistance have been identified. PROCEEDINGS OF THE LINNEAN Society oF New SoutH WaAtEs, Vol. 91, Part 3: 190 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS LITERATURE REVIEW Briggs and his coworkers at Davis, California, using a single race T-1 of of T. caries, have found the Martin gene, M, in nine varieties (Briggs, 1926, 1930b, 1931, 1932a, 1934; Smeltzer, 1952; Schaller and Briggs, 1955). Briggs (1926) showed that the variety Hussar possessed the Martin gene and a second major gene, H. From a cross between Hussar and the susceptible variety Hard Federation, the H factor was isolated in Selection 1403 (Briggs, 1930a). The Turkey gene, T, has been identified in six varieties (Briggs, 19320, 1934, 1936; El Khishen and Briggs, 1945). Analysis of all data accumulated from different crosses between varieties possessing the M and those the T gene indicated that the ratios deviated significantly from those expected on the basis of independent segregation. Briggs (1940) estimated the recombination value between the M and T genes as 34-22 units. A third major gene, R, was identified in two varieties (Stanford, 1941; El Khishen and Briggs, 1945). Stanford concluded that the R and T genes were closely linked with an estimated crossover value of 2-4 map units. From the tester cross results with Martin, the R gene was postulated to lie between the T and M genes. Schaller and Briggs (1955) presented evidence from their own results and from an analysis of combined data of certain previous publications which showed linkage between the H and M genes with a recombination value of 37-2%. The gene order HMRT for the linkage group concerned was established. The H gene would therefore be more than 50 crossover units from the R and T factors. Four weak resistance genes allowing moderate levels of infection in the homozygous state have been reported. The gene X, in addition to the R and T genes, was identified in a strain of Turkey wheat, C.I. 10016 (El Khishen and Briggs, 1945). This gene accounted for the spreading effect and absence of a definite mode in the susceptible group of F, rows in a cross between Turkey C.I. 10016 and the susceptible variety Baart and permitted about 25°% of bunted plants when homozygous. El Khishen and Briggs from studies on inheritance of resistance in a related Turkey strain C.I. 10015, concluded that this variety possessed only weak genes —X, and a second factor Y, which when homozygous was postulated to permit about 45°% of bunted plants. A deficiency of susceptible rows in tester crosses with Martin, Rio and Turkey, indicated that the Y gene was linked with the HMRT linkage group. Smeltzer (1952) established that bunt resistance in the variety Minturki was governed by two weak genes, U and V. The gene U permitted approximately 42 -5° infection in the homozygous state and 55°, when heterozygous. Comparable values for the factor V were 35 and 50%. One of these appeared to be closely linked to R and T, but it could not be determined whether U or V was involved in linkage. MATERIALS AND METHODS Turkey C.I. 10095 and C.I. 10097 are two out of 12 bunt resistant selections isolated from Turkey Red or South Dakota No. 144 wheat, a hard red winter type, by Kiesselbach and Anderson (1930) at Lincoln, Nebraska. C.I. denotes the accession number of the Division of Cereal Crops and Diseases, Bureau of Plant Industry, Soils, and Agricultural Engineering, United States Department of Agriculture. Originally head selections were made by Kiesselbach and Anderson from bunt free plants in a special planting of the Turkey Red variety heavily inoculated with a collection of 7. foetida made at Lincoln. With this bunt collection they found an average of 0-6°% infected heads in the 10095 Strain and 0-2°% infection in Strain 10097 in a series of annual tests over a seven-year period. During these tests the original unselected Turkey Red averaged 30:7% infection with the same inoculum. Turkey C.I. 10015 and 10016 studied by El Khishen and Briggs (1945) were from the same source. Each of these two resistant varieties under investigation was crossed with the bunt susceptible variety Baart. They were also crossed with the resistant E. P. BAKER 191 varieties Martin, Turkey 3055, Rio and Selection 1403, which are testers for the M, T, R and H genes respectively, and with Turkey C.I. 10015, which is the tester for the X and Y genes. F, progenies, produced from non-infected F, plants, of the crosses with Baart were grown at Davis, California in duplicate rod rows each containing 80 seeds. Prior to sowing the seed was thoroughly blackened with chlamydo- Spores of a race designated as T—1 of 7’. caries from its behaviour on the standard differential set (Rodenhiser and Holton, 1937). This collection has been perpetuated and used at this station since 1919; F, progenies of the tester crosses were handled similarly except that they were grown in single rod rows. Inoculated F, populations and appropriately spaced parental checks, including rows of the susceptible variety Baart, were included in the nursery. The plants were pulled when nearly mature and classified as smutted or non-smutted, the former category including plants with any evidence of bunt infection. EXPERIMENTAL RESULTS The mean annual percentages of bunt-infected plants observed in parent varieties in the present investigations over a 14 year period at Davis, California, are shown in Table 1. The mean percentage of bunted plants for each variety in all rows in the particular year in which inheritance studies were conducted is also indicated in the Table. Both Turkey C.I. 10095 (henceforth referred to as Turkey 10095) and Turkey C.I. 10097 (henceforth referred to as Turkey 10097) were highly resistant, the highest annual infections being 1-0 and 2-8% respectively. Both strains were consistently more resistant than Turkey C.I. 10015 (henceforth referred to as Turkey 10015), another strain from the same source. Generally Turkey 10095 showed a slightly higher level of resistance throughout comparative tests. TABLE | Percentages of Bunt Infection in Parent Varieties Mean of all rows in season Mean of 14 of current investigations Variety annual tests Turkey 10095 Turkey 10097 studies studies Baart . . J: Teel 79-0 82-8 Selection 1403 0-3 0-0 0-0 Turkey 3055. . 0-9 1:8 12 Rio... re 0-6 193} Iz) Martin a 0-0 0-0 0-4 Turkey 10015 3°8 4-1 ices Turkey 10095 0-3 0-2 — Turkey 10097 0-6 — ihodl The F, data for all crosses involving both Turkey 10095 and 10097, together with those for parental rows associated with the F, populations, are summarized in Table 2. As some genetically susceptible plants always escape infection in such tests and some genetically resistant plants may become infected as shown by Briggs (1926, 1929), F, data alone provide inadequately critical information on inheritance. For more accurate genetic analyses F, lines were used. The F, information, however, does serve to indicate approximately the number of genes involved in the resistant varieties under investigation, especially in the case of major or strong genes, the effect and mode of inheritance of such genes, and also to predict the behaviour of the heterozygous class in the F, populations. 192 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS TABLE 2 Percentages of Bunted Plants in the Parent and F, Rows of Crosses Involving Turkey 10095 and Turkey 10097 Number of plants Percentage of Parent or Cross a untpeaaplants Total Bunted ‘Turkey 10095 BG sip bie Be 1384 3 0-2 Baart . Pe a He Be 461 374 81-1 Selection 1403 op ae Be ae 119 0 0 Turkey 3055. ae ye ish BEA 242 3 1-2 ROME a ae or ws ar 233 4 1-7 Martin ane ale de Bis re 108 0 0 Turkey 10015 MG ne ax me 129 4 3-1 Turkey 10095 x Baart ; fs Lge 1482 394 26-6 ‘Turkey 10095 x Selection 1403.) op en 720 48 6-7 Turkey 10095 x Turkey 3055 aig a 707 2 0-3 ‘Turkey 10095 x Rio : ae =f 570 0 0 Turkey 10095x Martin... oh 5 692 10 1-5 ‘Turkey 10095x Turkey 10015... fs 772 0 0 ‘Turkey 10097 5 be oe ae 1425 17 1-2 Baart . Be Ne He a5 742 621 83-7 Selection 1403 bi oe or: Ae 128 0 0 Turkey 3055. ay as aii Be 202 4 2-0 Rio .. es is ae ae Bie 174 3 1-7 Martin ry oh ov, bs = 78 0 0 Turkey 10015 se off 3b a 92 9 9-8 ‘Turkey 10097 x Baart sit the i, 832 243 29-2 Turkey 10097 x Selection 1403... Si 447 4] 9-2 ‘Turkey 10097 x Turkey 3055 A a 528 25 4-7 Turkey 10097 x Rio ! ne a 594 10 Io} ‘Turkey 10097 Martin .. at ie 523 19 3°6 Turkey 10097 Turkey 10015... 10 306 19 5:9 The distributions of F, rows of crosses involving Turkey 10095 and 10097 are tabulated in Table 3 together with the distributions of the appropriate parental rows associated with the F, progenies. These distributions are shown by 5% classes with a separate classification of completely immune lines into a 0% class. Two weak factors, X and Y, additive in effect, were postulated by El Khishen and Briggs (1945) to be responsible for the resistance shown by Turkey 10015. A related strain Turkey C.I. 10016 (henceforth referred to as Turkey 10016) was considered by them to posses the factor X, in addition to the major R and T genes. As will be shown subsequently the presence of weak factors in both Turkey 10095 and 10097 was inferred from segregation in crosses with tester varieties but primarily from the pattern of distribution of F, rows in their crosses with the susceptible variety Baart. Weak or minor genes cannot be distinguished and documented with the same precision aS major genes unless they are each first isolated into separate lines, which then can be intercrossed. On the basis of inheritance studies conducted in this manner, a decision then may be possible as to whether or not each gene is different and distinct. Obviously until such procedures are adopted a minimum number of minor genes should be invoked which will satisfy and reconcile segregation patterns exhibited in all crosses involving varieties where weak genes are considered to be operative. This principle was applied in the case of the four Turkey selections, two in the current investigations, all from the same source studied at the California station. 193 EH. P. BAKER SMOI JO riequinu 1210.1 G g I I G v G I I I v (s} I 6 6 LT I 6 I G I L v 8 Or OL v 8 L 9 8 St OL 1 OG Ge OF GI O¢ S&T T g L I I I G I G G I T G I I I I G I I I G I G L g G 9 v 9 9 L 9 € 9 sit GI OSE SE BIE = UL I te g te I OOT— §6-— O6- 8 O8 GL OL G9- O9- GE— OS— GH OF GE- OF GE-— O06— No) i) SNE SI) Iv OI- v ‘pepnpour Ajuo sMoa oyeortdnp jo weep, 7 GIOOT Reeatcuiwny Aoyrn J, ulgIeA X L6001 Aoymy, " Ore xX 16001 Aeqmny, "* gcgog Aoyamny, X L6001 AexmM I, is E0FL Woroefeg X L6001 Aoxrny, _reeg x 146001 ey "* gT00T Aoymy, % ae ca . - =f a0 o "* geog Loyang, Sc OVienonaeles g * qaeeg i Se :600 Tees ; “SIOOT ‘sexamiy, X $6001 Aoymn J, urqaeT X 6001 Aoymy, " ora X g6001 Lexi, "* Geog sowang, X 46001 Aoymny, GOFT WoryoeTeg <¢600T Aoxmny, reed X 96001 Loy “* gTo0T Aoymy, "Ula, = =a * art a "* geog Aoymy, 7 ‘ €0FI Wooeeg ¢ “+ gree c600T Aoxany, G— 1°96 1:06 T-G8 1-08 I-92 1-02 T-99 T:09 I-99 1-09 I-:Sh 1:07 1:GE 1-0€ 1:SG 1:0G 1:ST T:Ol I-g T:0 O sessejo yuoorod g Aq sMOd Jo WOTyNqIaASTCG SsOd) 10 JUOIe uoyoafuy qung sof sassniQ qwaoleg G one LEQ0T haying, puo GEQOT hayimy, buaajoauy sassoug fo snoy *q pun ojuamg fo uoyngysrg § WiIavy,; 194 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS To implement this objective some minor modifications in the analyses of Turkey 10015 and 10016 as proposed by El Khishen and Briggs (1945) were considered necessary. The evidence for these and the steps which were followed in arriving at appropriate modified genotypes for these Turkey strains were as follows : (a) Although transgressive segregation for bunt susceptibility was recorded in the current investigations in the cross Turkey 10097 xTurkey 10015 no susceptible F, rows were obtained. However some F, rows showed 20-25% infection, which suggests that these varieties have in common the stronger, or X, gene of the two (X and Y) reported by El Khishen and Briggs in 10015, since the factor X, when homozygous, was considered to permit 22-5°% infection. (b) In the case of both Turkey 10015 and 10097, segregation producing susceptible F, lines occurred as expected in their crosses with Selection 1403 on the hypothesis that the H gene in Selection 1403 is more than 50 crossover units from the factors for resistance in these Turkey strains. In the case of the genetic testers for the M, R and T genes (which have previously been demonstrated to be genetically linked) a consistent deficiency of susceptible F, progenies was evident on the assumption of independent gene segregation in crosses involving these testers with both Turkey 10015 and 10097. The segregation pattern observed in both strains was parallel with each tester cross as a comparison of Table 3 in the present publication and Table 3 in the paper of El Khishen and Briggs shows, allowing for the fact that the level of infection in the susceptible Baart check rows was some 10 to 17% higher in their tests. It is probable, therefore, that if both Turkey 10015 and 10097 have the gene X in common, this factor is linked with the HMRT linkage group. The absence of moderately susceptible rows in the cases of Rio (R) and Turkey (T) tester crosses with Turkey 10097, and in the case of the Turkey (T) tester cross with Turkey 10015, further indicates a close linkage between the X with the R and T genes respectively. Unless this close linkage is assumed a reasonably high proportion of F, progenies heterozygous for X and lacking other genes for resistance would be expected; these lines would, on the degree of infection postulated for the heterozygous Xx condition, be expected to show approximately 60% infection. No F, row with such a high infection percentage was actually observed in these crosses. (c) That some slight revision would be required in the genetic analysis of Turkey 10016 as published by El Khishen and Briggs is indicated from these observations. The present hypothesis which slightly modifies their analysis is that Turkey 10016 has both the X and Y genes in addition to the major R and T genes, with X, as previously indicated, and not Y being in the HMRT linkage group. Although it is realized that the same precision in genotype analysis where minor factors are implicated cannot be expected from the degree of Segregation observed in tester crosses other experimental evidence supports the conclusion outlined for the genetic constitution of Turkey 10016. No F, rows showing more than 10% infection were reported by El Khishen and Briggs in the cross 10015 x 10016. On the basis of linkage between X and the R and T genes with 10°, crossing over between R and X (a value finally decided upon from analyses of Turkey 10015, 10016, 10095 and 10097 hybrids with tester varieties) 12 such rows would have been expected. On the postulated order for the M, R and T genes proposed by Stanford (1941), since X appears to be fairly closely linked with M as well as R, the X gene lies between M and R but closer to the latter, the gene order in the linkage group being thus MXRT. For all classes a crossover value of 10% between X and KR gave the best overall fit between observed and expected results for the cross Turkey 10016 x Baart and also for the tester cross of this variety with Martin. Values consistent with previously known information were assigned to F, rows derived from different F, genotypes and values which might reasonably E. P. BAKER 195 be expected on gene interaction assigned in other cases. The theoretical distribu- tion for the cross Turkey 10016 x Baart was calculated by the method to be indicated in the discussion under hybrids involving Turkey 10095. The +?,, value for agreement between observed and expected results was 6-83, giving a P value between 0:95-0:90. The revised genotype of Turkey 10016 was thus con- sidered to be XXRRTT YY. ANALYSIS OF GENOTYPE OF TURKEY 10095 IN RELATION TO BUNT RESISTANCE The distribution of the means of duplicate F, rows on the basis of 5°/ class intervals for the cross Turkey 10095 x Baart is shown in Figure 1. This curve is based on per cent of rows in each class out of the 225 rows actually tested. The per cent distribution was adopted so as to serve as a basis for comparison with previous similarly depicted distributions. It will be seen that there are two minimum points on the curve—at the midpoints 12:5% and 57-5% respectively. The curve has, in general, features similar to those published by Briggs (1936) for the distribution of F, rows in crosses involving certain Turkey varieties and Baart in which segregation was due to the action of a single incompletely dominant gene present in these Turkey varieties. The main differences are that the heterozygous class shows a more general spreading distribution with absence of a pronounced modal class, and that there is also a definite spreading effect with lack of a definite mode for the susceptible group. This latter effect indicates the segregation of one or more weak genes. Further- more, if 12-5 and 57-5% are taken as minima on the basis of the action of a single gene there are obviously too few susceptible rows for the single gene hypothesis to be statistically satisfactory. Again if the minima at 12-5 and 57-5% are taken to indicate the separation of the F, lines into a monohybrid 1:2:1 ratio, the infection percentage in the then heterozygous group of F, rows Should approximate the F, percentage infection. The mean per cent infection in the EF’, rows in the cross Turkey 10097 x Baart with between 12-5 and 57-5°% of bunted plants was 34:9, compared with a corresponding F, population infection of 26-6% as indicated in Table 2. This further suggests that the segregation observed was due to the effect of one or more weak genes as well as a major gene for resistance. Information as to the identity of these genes is obtained from an analysis of F, and F, data on crosses between Turkey 10095 and tester varieties. Segregation producing susceptible F, plants and F, rows was obtained in crosses involving Selection 1403, Turkey 3055 and Martin, indicating the H, T and M genes respectively were not present in Turkey 10095. However, the absence of any segregation in both the F, and F, generations of Rio and Turkey 10015 crosses with Turkey 10095 indicates that the strong gene is the Rio (or R) gene and that the weak gene (or genes) is one (or both) of those present in Turkey 10015 (X and/or Y). No susceptible F, plants or segregating F, rows were produced in the two crosses indicated. With R as the major gene for resistance the presence of one or more weak genes in addition is also necessary to account for the low F, infection of 26-6% in the cross with Baart. Stanford (1941) found in the F, of a Rio x Baart cross 45-6°% infected plants, which was con- firmed by the heterozygous F, lines averaging 51-4° infection. The presence of both X and Y as weak genes is favoured for the following reasons, the evidence being briefly summarized for each point : (a) More susceptibility in the F, and F, generations of the cross between Turkey 10095 and Turkey 10015 would be expected if only either X or Y was present. Computations show that with the stronger or X gene alone present, 1/16 of the F, plants of this cross would be homozygous XX in genotype, with this as the only gene for bunt resistance. The previous behaviour of this gene indicated that F, rows derived from such genotypes produce approximately 22-5% infected plants. Also certain other genotypes would be expected to 196 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS produce F, lines with a mean infection above 5%. Computations based on the presence of the Y gene alone with 10% crossing over between the R and X genes indicate that only 0-2% of F,; rows would possess the homozygous Y gene alone for resistance. However, 4:5° would be expected to be derived from plants of the genotype Rrxx YY with a predicted amount of infection based on factor interaction of approximately 25-0%. A similar percentage of rows would be derived from rrXx YY F, plants with a somewhat higher predicted F, infection percentage. Finally 40-5% of F, plants would possess the R and X genes in repulsion and calculations assess an infection of approxi- mately 10° in F, rows derived from such plants. The fact therefore that such a large proportion of F, plants theoretically produces F, lines which would be expected above the 10% infection level supports the hypothesis based on the presence of both X and Y as weak resistance genes. (b) Although behaviour resulting from the interaction of genes conditioning bunt resistance cannot be predicted it would not be expected that either X or Y in conjunction with the R gene would confer the low level of F, infection (26 -6%) observed in the cross Turkey 10095 x Baart, since Stanford (1941) has shown that the R gene when heterozygous permits about half the plants to become infected. (c) An analysis of the comparison between observed and expected results for F, data of the cross Turkey 10095 x Baart indicates that both the X and Y weak genes are present in the former variety. Assuming firstly that only the genes R and X are present in Turkey 10095 and linkage with a crossover value of 10% between them, then 56 F, rows would be expected above the 55% infection level from the two genotypes rrxx (85% similar to susceptible Baart) and rrXx (62-5°%,), whereas only 40 were actually observed. Of the 56 expected lines in this category, 46 would be considered to have approximately 85% infection, whereas only 24 were observed above 70% infection. A comparison likewise between observed and expected results for the distribution of F, rows was made on the assumption that the independent R and Y genes alone were present in Turkey 10095. This comparison was entirely satisfactory in the resistant classes and also those showing above 55%, infection, the calculated figure for the former group being 56-7 and the latter 46-7, compared with observed numbers of 68 and 40 respectively. However, if the F, infection of 26-6% is used in predicting the behaviour of the doubly heterozygous class in the F, the genotype Rr YY would be expected to be slightly more resistant than this. When the genotypes Rr Yy and Rr YY (3/8 of the population) are normally distributed on this basis, the observed 30-35% classes have a corresponding surplus. The y?,, value for all infection classes was 39-86, with a corresponding P value of <0-01. For these reasons the genotype of Turkey 10095 was considered to be RRXX YY with regard to genes conditioning bunt resistance. On this hypothesis a comparison was made between observed and expected results for the distribution of the 225 F, duplicate rows of the cross Turkey 10095 x Baart. The 10° crossover value between R and X previously postulated for Turkey 10016 in the introduction to this Section gave a satisfactory fit and the analysis in the cross with Baart and also with the tester varieties was not improved by varying the crossover value between them. The Y gene was considered independent of the other two. That rather close linkage exists between K and one of the weak genes was also shown by a comparison of observed and expected results with independent behaviour of these three genes. Under this latter condition only 17-5 of the 225 F, rows would be expected to exhibit above 55% infection which is in serious disagreement with the 40 rows actually observed in this category. Linkage with 10% crossing over between R and X was finally adopted then, since, considering the four varieties postulated to contain the X gene—Turkey 10015, Turkey 10016, Turkey 10095 and Turkey E. P. BAKER 197 10097—best agreement was obtained between observed and expected results using this value. On this basis the proportion of F, genotypes, the number of F, rows expected from each out of the total of 225 and the per cent of bunt expected in F, rows from the different genotypes are indicated in Table 4. The values assigned to the different genotypes in general were very close to those previously reported. The mean of the Baart checks sown throughout the F, and F, rows in the cross Turkey 10095 x Baart was 81-9 with a standard error of 1:35°%. The mean+2-18 times (the t value at the 0-05 probability level for 12 degrees of freedom) the standard error is therefore 81-9-+2-94 or 84-34%, with 85 as the closest integer. The value of 85° thus used for the 1rxx yy genotype, which would be expected to be similar to Baart, did not therefore differ significantly from the 81-:9°% mean for Baart and gave a better agreement than the actual Baart mean for the most susceptible class; F, rows produced from different genotypes involving X and Y without the R gene were assigned values by El Khishen and Briggs (1945), and these values were followed in general in assigning values to the F, rows derived from such genotypes, except that the values for Xx yy and xx Yy were approximately 10% lower than assigned by these authors. The mean of the Baart checks to which their data refer was 93-5°%, however, compared with 79-0°% in the present investigation. It is logical therefore to lower the infection level of the genotypes Xx yy and xx Yy which confer little resistance on F, lines accordingly by a somewhat less amount than the difference in the Baart checks in the two years under comparison. The values for the other genotypes involving X and Y without R were identical to or within 2-5°%% of those postulated by El Khishen and Briggs. The genotype containing R in the heterozygous condition alone was ascribed a value of 50-0, a figure suggested by Stanford (1941). Information on the F, behaviour of the triple heterozygote was obtained from F, data. The mean infection in the F, was 26:6%. A much better agreement between observed and expected results was secured by assigning a higher value to rows derived from this genotype in the F,. The standard error J od of the F, based on 1,482 plants was 1-:15°%. The upper 95% fiducial limit was 28-85% and a value of 29-0°% (the nearest integer) was assigned to the F, triple heterozygote. The most complete agreement between observed and expected results was secured by giving this genotype a value of around 35%, but this would have differed significantly from the 26-6% observed. No attempt was made to assign values to the different F, rows indicated as resistant although it is recognized that these would differ slightly in resistance. Any genotypes containing R in the homozygous condition would be expected to show a mean not exceeding that of Rio (1-3%), whilst those containing the XX YY genotype (2-66 rows) would exhibit a mean of 4-:1% similar to Turkey 10015. On a binomial distribution for these mean percentages all would be expected to fall below 10° infection and the resistant genotypes were arbitrarily assigned to the 0-5 and 5-10 per cent classes With regard to the other genotypes although no positive evidence is available on gene interaction, values were assigned which seemed logical on the behaviour of these genes independently. Except for the triple heterozygote the various genotypes permitting bunt were assigned mean values which were multiples of 2:-5°%. Further refinement would have resulted in a closer agreement between observed and expected results, but this did not seem justified as it would have added little further significant information in the present instance. A difference in reaction of genotypes with the R and X genes in the coupling and repulsion phases respectively would be expected and is indicated in Table 4. 198 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS Frequency distributions were calculated for all genotypes other than those classified as resistant in Table 4. For means above 10% this was done by using the means indicated and the standard errors for each mean derived from the formula, s= (| ~s where p and q are the infection percentages expressed pro- portionately, and n is the mean number of plants per duplicate row or BESS TABLE 4 Proportion of F, Genotypes Expected, Number of F; Rows Expected out of a Total of 225, and Percent of Bunt Expected in Ff, Rows of Turkey 10095 x Baart (Based on linkage between R and X genes with 10% crossing over) F, genotype Number Mean F, genotype Number Mean and of percent of and of percent of proportion F, rows bunt expected proportion F, rows bunt expected expected expected in F, rows expected expected in F, rows RX Y i TONG NYS RX Y 0: 0506 11-39 Resistant Rs ¥ 0:0013 0:29 12-5 eS Genie 22-79 Resistant TX Y 9.0025 0-56 20-0 ie WW 5 Rx Y as : 0-0506 11-39 Resistant = 4 0-0013 0-29 27-5 Ike NL : exe WY RX 7 0:0112 2-5 Resistant Rx ¥ 0-0112 52 25-0 Rx y ? rx y . : RX ¥ 0:-0225 5-06 Resistant Rs Y 0:-0225 5:06 42-5 Rx y ‘ : eS. 7. oN ay RX 7 0:0112 2-52 Resistant Rx = 0:0112 2-52 50:0 rx Y F rx Y : RX ¥ 0-0112 2-52 Resistant xX ¥ 0: 0006 0:14 Resistant RI 3 : rX Y 9. a) . RX ¥ 0:0225 5-06 7:5 X Y¥ 0:0013 0:29 12-5 PIX rm y RX = 0:0112 2-52 10-0 x = 0- 0006 0-14 22-5 ws VW ip YY eles ashes ; ‘ ney Sos . 9. 3 RX Y 0:-1013 22-79 20-0 xX ¥ 0-0112 52 34-0 RX Y 0-2025 45-56 29-0 xX ¥ 0:0225 5:06 52-5 rx y TEXTS) 9. Ke aS YL Molla 2 : y , -52 : RX y 0:1013 22-79 40-0 x oe 0-0112 2-5 62-5 Rx Y ¢ rx Y Rx ¥ 0-0006 0-14 Resistant = ¥ 0: 0506 11-39 45-0 Rx Y 9.0013 0-29 Resistant = oon 22-79 70-0 Ieipze NS x Y R ~~ ¥ 0-0006 0-14 Resistant ~*~ YZ 60-0506 11-39 85-0 Rx y tmx y 199 H. P. BAKER GO-00L T0-00L 3% 68:23 06:86 OF-IT 08:2 8E-IT G4-G 80-G O0G:G GI-0 82:0 09-6 90-G GG-G 66:0 99:0 86:0 84:26 99:SF 84-26 2G-G 90-S PIO. 00-0 10-0 0 60:0 60:0 0OT—-S6 68-0 Gé-0 G 84-0 84-0 G6 —06 19-6 81°36 9 16-7 06-7 10-0 06 —S8 84-1 66°% i TL: 06:F 2-0 G8 —08 L19°@ Tg-T i) 66° 84-0 09°46 10-0 08 —GZ 19-6 98-6 9 69-8 20:0 49-8 GL-0 GZ -02 IL-& 60:7 Z 61-6 Gg. 8 19:0 €0:0 04 —99 19°@ 92-1 9 G6-& 09:6 10:0 ¥0-T 92-0 t0:0 G9 -09 66-1 9T-T & 19-G %2:0 81-0 19-0 &@-T Gé-0 c0-0 10-0 09 —S¢G 19-G L1E-%G 9 IL-G 10:0 GP-T Gt-0 0-6 68-0 926-0 PE-0 CASE 0} 4) 19-9 LG-P GI 86:01 GO:F 10-0 €@-T &0-0 68:0 G6-T 98°36 0S —SP tP-8 98-9 61 TPS GO: 96-0 0&-0 G&:0 90-6 61-8 &2:-0 Gy —0P 68-8 8-9 06 Gé-ST GPT 60:0 6-0 70:0 Gé-T 20-0 10-0 61-8 L7-& OF —S& 19-9 89-8 GT TG-6L 8L-0 66-0 96:0 96:0 20:0 G8-G 88-FL 60-0 Gé —0& 00-8 9T-OL 81 98-636 10-0 0€-0 60-0 c0:0 86:0 €1:0 40-0 TE-0 96-8T £0-G 0& —GG 82-G G6-2 &L 88°21 60:0 90:0 °c0-0 86:0 240-0 €2:0 20-0 10:0 61:4 26-6 GE —0G 68-7 68-7 IL 68-01 60-0 70-0 96:0 10-0 €2:0 ¥F0-0 08:0 46:6 &1:0 20-0 06 —SL 19-6 68-1 9 OL-F 9T-0 60:0 GO-:0 91-0 60:0 60-6 66:0 99-0 GT —OT 07-8 64:6 61 68-16 G2-9T F0-0 t0-0 60:0 &&-T F9-& OL —G IT-6L GL-6T 8 &F 00-h GI-GP 60:0 60-0 40-0 4:0 & —O0 (gU9010 J) (qU90IO J) (TOqQUIN jy) (LOquUAN Ny) (ToquuN jy) SMOL SMOL SMOI SMOI SMOL z 7 ra 7 - . 4 . a 7 a ssvpo pearss pojood poares pojood yurysis £ = 4 SS Au tx a se AXL AME AM pec ee ee eal a a ot pe WOTOoJUI -40 -XOP -40 -XOD -o-f RR sat MN sai He Mah AR see AR orate 7N Sean AN at AN AN eae IN Sa eae AN xa AN par AN aE NK aE ON OK aK dE WN Rcd AN 38a qung, (F e[qeyz, UL poxyeorput sedAqoues I0y posn suvoUT puv SMOAI JO JOqUINN)) paywoupuy sadhjouay josasnag oy. sof wMVg Xx CEOOL Kayan fo smoay *y fo suorungiysig 1n91YW2eL00Y 7, G HIaVy, 900 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS plants in the present instance. The proportion of each genotype which lay in the successive class intervals around the mean was then calculated on the basis of a normal distribution from the ratio Moreen . Distributions with standard error means of 7-5 and 10% were calculated on the basis of a binomial distribution in each case according to the formula (p+q)®. Comparisons showed that with regard to means above 10% the normal distribution very closely approached the binomial distribution and was accordingly used. These frequency distribu- tions account solely for sampling errors and probably underestimate the errors involved in distributing the rows of each genotype since no account is taken of environmental effects. The separate distributions of the genotypes concerned are shown in Table 5, as well as the total number of rows expected in each 5% infection class, by taking the combined values of the separate distributions. The comparison between observed and expected distributions is shown in Figure 1, indicating the per cent of rows in each infection class. The chi-square test for comparison of observed and expected number of F, rows in each class is presented in Table 6. In this test the resistant rows in the 2-5% and 7-5% classes were grouped together as the expected rows had been assigned in this way in these classes. Classes with expected numbers less than five were grouped with an adjacent class or classes to give an expected number greater than five. The total y?,, value was 11-8205, giving a probability value between 0-70 and 0-50. The proposed genotype for Turkey 10095 will now be considered in a comparison of observed and expected results in tester crosses other than those involving Rio and Turkey 10015, which have previously been considered. In calculating expected distributions in these instances the order of the linked genes was taken as Turkey (T), Rio (R), X and Martin (M) in conformity with the relationship suggested by Stanford (1941) in assessing the linkage between the M, R and T genes. The map distances T-R, R-X and X-M on the basis of 10% crossing over between R and X were taken to be 2-4, 10 and 20 crossover units respectively. These figures, especially for the X gene, are considered to be only approximate but are in close agreement with the figure of 34-22% given for the linkage between the Martin and Turkey genes (Briggs, 1940) and the estimate of 2-4°% linkage between the Rio and Turkey genes made by Stanford. In any case, the map distance between M and T should perhaps be increased as the intervening genes allow the detection of double crossing in this region, but further refinement does not seem necessary concerning this point at this stage in view of the uncertainty of the extent of linkage between R and X. As indicated in Tables 2 and 3 segregation occurred in tester crosses with Selection 1403, Turkey and Martin, indicating that neither the H, T or M genes respectively were present in Turkey 10095. In the cross with Selection 1403, two F, rows were observed with above 60° infection whilst computations show that 4:08 would be expected in this category, derived from the three genotypes —completely susceptible, Xx and Yy. Although the gene H is on the same chromosome as X, they are probably more than 50 crossover units apart, on the data of Schaller and Briggs (1955), and are therefore genetically independent. A binomial distribution calculation shows that the probability of obtaining a deviation as great as or greater than that observed through chance alone is 0-34. A chi-square test was not used as one of the expected classes was less than five. As expected, less segregation occurred in the crosses involving the T and M genes on account of linkage with R and X. In the test cross with Martin 1 row above 60% was recorded and three above 45%, the computed values being 0:78 and 1-78 respectively ; obviously the agreement between observed and expected results in the former case is entirely satisfactory, and on the basis of a binomial distribution the probability of obtaining a deviation as great as or greater than that observed through random sampling was 0-43 E. P. BAKER TABLE 6 x2 Test for Comparison of Observed and Expected Number of Ff; Rows for the Cross Turkey 10095 x Baart, for the Several Bunt Infection Classes Indicated 201 Bunt infection Number observed Number expected Deviation x? class Rows Rows oO 5 43 43-00 : 5— 10 19 a0 f spay O88 10— 15 6 4-10 15- 20 ib 10-83 f ay Ue2t50U 20— 25 13 17-88 —4-88 1-3319 25— 30 18 22-85 —4-85 1-0294 30— 35 15 19-51 —4-51 1-0425 35— 40 20 15-35 4°65 1-4086 40— 45 19 15-41 3°59 0-8363 45— 50 15 10-28 4-72 2-1672 50— 55 6 5-11 0-89 0-1550 55— 60 3 2-61 ‘ : 60_ 65 6 3-95 f 2-44 0:9076 65— 70 7 9-19 —2-19 0-5219 70— 75 6 8-69 —2-69 0-8327 75— 80 6 3°39 4 : 80— 85 a Roi 1-47 0- 2533 85— 90 6 4-91 90— 95 2 0:78 2-29 0:9184 95-100 0 0-02 Total 225 224-89 11-8205! ne Number of rows, per cent. as He aaa aca Ne2ee ZIT 2D) AT IZENLLD 32,0 379 429) 47-9) S27) 62 SOF 9) i ie 82:5 875 925 975 Bunt Infection, per cent. Fig. 1—Observed percentage distribution of F; rows of the cross Turkey 10095 x Baart for bunt infection (solid line) compared with theoretical distribution (broken line). 902 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS in the second comparison. With the tester cross involving Turkey 3055 the close linkage between the T, R and X genes gives a low probability of F; rows above 60% infection being produced, only one row in about 15,000 being computed to fall into this infection range. However above 40% 0-69 rows out of 250 tested derived from various genotypes—completely susceptible, Xx, Yy, Xx Yy, Tt, Rr, YY and Rr Yy, which would reasonably be expected to fall above this infection point—would be theoretically expected, whilst as shown in Table 4 one was actually observed in the 55-60 infection class. The postulated genotype for Turkey 10095, RRXX YY, with 10% crossing over between the R and X genes, with regard to bunt resistance therefore gives satisfactory statistical agreement between observed and expected results in crosses both with Baart and tester varieties of known genetic constitution for bunt resistance. ANALYSIS OF GENOTYPE OF TURKEY 10097 IN RELATION TO BUNT RESISTANCE An analysis of the genotype of Turkey 10097 indicated that its resistance was due to the additive or complementary effect of two weak genes, one being the X factor and the second a previously unidentified factor, designated W. The distribution of the per cent of F, rows in the cross Turkey 10097 x Baart is shown in Figure 2. The presence of two weak genes in Turkey 10097 was necessary to account for the F, infection of 29% in its cross with Baart and for the spread of F, rows in the same cross. With the X gene alone, the predicted behaviour would give an F, infection of approximately 60°% with a corresponding grouping of F, rows in the intermediate class around this mean, with one-fourth of the rows, comprising the completely susceptible group, above this figure. As indicated in Table 2 and Figure 2 this hypothesis falls far short of fitting the observed results. The absence of a maximum point in the group of resistant F, classes indicates that the factor W must confer little resistance even in the homozygous condition. TABLE 7 Ratio of F, Genotype, Number of Ff, Rows, and Mean Percent of Bunt in F, Rows Postulated for the Cross Turkey 10097 x Baart, the X and W Factors Being Independent Ratio of F, Genotype Number of F,; Rows Mean Percent of Bunt in F,; Rows 1 XX WW 15-19 1-0 2 XX Ww 30-38 9-0 2 Xx WW 30-38 20-0 4 Xx Ww 60-75 32-5 1 XX ww 15-19 22°5 2 Xx ww 30°38 60-0 1 xx WW 15-19 37-5 2 xx Ww 30-38 47-5 1 xx ww 15-19 80-0 Total 243-03 The X and W genes were considered independent and the values assigned the mean percentage of bunt expected in the F, rows in the various genotypes ‘in the cross Turkey 10097 x Baart are indicated in Table 7, together with the number of F’, rows expected for each genotype. Of the various values assigned the W gene, those which gave the best agreement between observed and expected results indicated a value of 40% infection in F, rows for homozygous WW and 50% for the heterozygous Ww condition. The values assigned XX and Xx were 22-5 and 60-0% respectively, compared with corresponding values of 203 E. P. BAKER 66°66 66°66 EPG 66°GPG G6'GI 96-0 61:-ST 9€-0€ 61-°STI GL-09 96-08 LE&-0& 61:ST [BI0.L 00-0 00-0 0 10:0 10:0 OO1-S6 00-0 60-0 0) 1é:0 1¢:0 C6 —06 00-0 LL-O 0) 98-1 98-T 06 —S8 G9°T 86° a7 PG-G €G-¢ 10:0 G8 —08 66° GEG 8 69°¢ 6¢-¢ OL-0 08 —SL 88°G 8i-T L 98° 98-1 00-1 GL —-OL LE:G c6-1 9 GL: 1¢:0 60:0 I¢:-P OL —S9 66:6 OL: F 8 96:6 10:0 66-0 99-6 G9 —09 L1:9 66°F GI 66°11 GV'G 10:0 9¢-6 09 —g¢ GIL‘? 68°F OL 68-11 SI:L LI-O Io? 60-0 Gg —0¢ 9L-G Gé-s Tl 10:°€1 G&: OL FI-1 00:1 G¢c:-0 OG —GF 66:8 vG:9 0G 91:S1 SI:L 69°& Or-0 96°F GP —OP G0-6 1:6 GG 8°66 GV'G 1€-¢ 10:0 90:0 LG: FI 10:0 OF —SE GE-Gél 8I-I1 O0€ 91: LG 66:0 69°S 18:0 66°16 1?:0 Ge —0& LT:9 6S:-6 GT €1:&¢6 60:0 PI-T 89°& LG- FI IL-€ O€ —GEé €¢6°8 68-8 0G 19:16 LI-0 60:9 96:7 GO-11 ¥0:0 GE —-06G L1-9 08:9 at 6-91 10-0 89° cc-0 GO-I1 vel 0G -SI LI-9 Gé:9 GI GL:SI 18-0 €0-°0 Wok LG- OT cI —O1 Iv: L 66:9 81 18-91 90-0 I?:0 66:91 éL:0 OL —& 8¢S-9 GI-L 9T 8é-LI 10:0 O€-G LO-ST G = (qued18q) (queo10q) (aoquin yy) (aequin yy) MM XX MAA XX AAM XX MM XY MM XK MAM XK MAXX MM XK MM XX SSBjo SMOI SMOL SMOL SMOL UWO1}OeFUL pearosqeo poyoodxa pearosqeo poqoodxa edAyouer) yung (L 9[qBjz, UI pozvorput ore sodAjoues Toy posn suvey() paqwoupuy sadfjouasy jo1eaag ay, of wong xX L600T Rayan fo sno * 7 fo suoynguysiq jvo4ye10eY [, 8 WIav 204 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS 92-5 and 62:5% in the case of Turkey 10095. Tests involving the two Selections were conducted in different seasons, the corresponding completely susceptible genotypes being assigned values of 80 and 85% respectively. Hence some slight reduction in the value assigned the heterozygous Xx condition might reasonably be expected in the cross Turkey 10097 x Baart. The mean infection in the 15 Baart checks throughout the F, and F, rows of the cross was 83-8-+1-49%. The lower 95° fiducial limit was 80-6% and a value of 80% for the completely susceptible xx ww genotype was adopted rather than the mean for the Baart checks as better agreement between observed - and expected results was secured. Similarly in the case of the double hetero- zygous genotype the upper 95% fiducial limit value of 32-3% for the F, mean of 29-2+1-58%, was used in distributing F, rows from this genotype. The double homozygous resistant genotype was assigned a value of 1-0°% similar to the mean of Turkey 10097. The two remaining genotypes XX Ww and Xx WW were assigned values which might reasonably be expected from the interaction of the genes concerned. The theoretical distributions of F, rows in the various genotypes were calculated as in the case of Turkey 10095. The theoretical distribution of the F, rows for each genotype is shown in Table 8, together with the computed total number for each infection class. The chi-square test for comparison of observed and expected numbers of F, rows for the different infection classes is shown in Table 9, the P value being 0-70-0-50. The theoretical distribution is compared graphically with the observed distribution in Figure 2. The geno- type XX WW for Turkey 10097 thus gives satisfactory agreement between observed and expected results for the distribution of F, rows of the cross Turkey 10097 x Baart, when the indicated values are assigned to F, genotypes. The postulated genotype for Turkey 10097 will be considered in its behaviour in tester crosses in greater detail. In the cross Turkey 10097 x Selection 1403, 3/64 of the F, population, derived from the genotype hhxx ww (completely susceptible) and hhXx ww (60% infection) would be expected to exhibit more than 55°% infection. In 104 rows tested four such rows were observed compared with 5-07 expected. On the basis of a chi-square test this comparison gives a P value between 0-70-0-50. With 20% crossing over between X and M, 2-25% of the F, rows in the Martin cross, derived from similar genotypes to those above, would be expected above the 55% infection class. In 106 rows observed, one was found in the 50-55% infection class. If this is considered to belong to the category above 55%, the probability on a standardized normal variate test of such a deviation being due to chance alone is 0:52. In the case of the cross involving Turkey 10015, as previously indicated, the infection range with the X gene in common would be expected to extend up to 20-25% in F, rows and two rows were actually observed in this infection class. The distribu- tion of infection in F, rows around a mean of approximately 10°, substantiates the theory that independent segregation of the W and Y genes occurred in this cross with the X gene present in both varieties. A comparison of observed and expected results in the case of tester crosses with Turkey 3055 and Rio was not entirely satisfactory however. Assuming the same gene order for the T, R, X and M genes as employed in the case of. the Turkey 10095 tester crosses, 3-5 F', rows of the 251 tested in the cross with Turkey 3055 would theoretically exhibit approximately 60°, infected plants. No rows showing more than 30-35% infection were recorded. Similarly three F, rows in the tester cross with Rio would be expected to be derived from F, plants heterozygous for the X gene alone, theoretically showing 60% infected plants in such lines. One segregate was recorded in each of the 35-40 and 40—45°% infection classes. However the fact that no moderately susceptible F, lines corresponding to those expected from appropriate genotypes in the case of both the Turkey BE. P. BAKER 205 TABLE 9 x2 Test for Comparison of Observed and Hxpecied Number of F, Rows for the Cross Turkey 10097 x Baart for the Several Bunt Infection Classes Indicated Bunt infection Number observed Number expected Deviation x? class Rows Rows é 0O-— 5 16 17-38 —1-38 0- 1096 5— 10 18 16-81 1-19 0-0842 10— 15 15 15-12 —0-12 0-0010 15— 20 15 16-53 —1-53 0-1416 20— 25 20 21-61 -1-61 0-1199 25— 30 15 23°13 —8-13 2-8576 30— 35 30 27-16 2°84 0-2970 35— 40 22 22-38 —0-38 0-0065 40— 45 20 15-16 4-84 1-5452 45— 50 14 13-01 0-99 0-0753 50— 55 10 11-89 —1-89 0-3004 55— 60 15 11-99 3°01 0-7556 60— 65 8 HE =1-96 0:3857 65— 70 6 4-75 70— 75 7 2-86 f 5:39 3:8176 75— 80 8 5:65 2°37 0-9977 80— 85 4 Dee | 85— 90 0 1-86 : 90_ 95 0 0-21 F —3- 62 1-7197 95-100 0 0-01 } Total 243 243-00 13-2146! 1D.f.=15, P=0-70—-0-50. Number of rows, Pel: cent. 25 75 125 175 225 275 325 375 425 475 525 575 625 675 725 775 825 8&5 925 G75 Bunt Infection, per cent. Fig. 2.—Observed percentage distribution of F, rows of the cross Turkey 10097 x Baart for bunt infection (solid line) compared with theoretical distribution (broken line). 906 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS 3055 and Rio tester crosses was not statistically significant at the 0-05 probability level on the basis of a standardized normal variate test. In any ease all tester crosses involving Turkey 10097 were sown in a section of the nursery where low bunt infection was recorded in the Baart check rows. Under these circumstances the infection in segregating rows would also be expected to be lower. Hence the two weak genetic factor genotype (XX WW) for Turkey 10097 gave satisfactory agreement between observed and computed distributions also in tester crosses, and this genotype is therefore proposed for this Turkey Strain. DISCUSSION Weak genes for bunt resistance have been reported by El Khishen and Briggs (1945), Smeltzer (1952) and in the current investigations. These are genes which are presumably incompletely dominant in all instances where they have been postulated to operate and which when homozygous permit up to 45°, of plants (in the case of the Y gene) to become infected with bunt. The gene X in this category was found to be associated in the present studies with the HMRT linkage group and was presumably located between the M and R genes but closer to R, with which it showed approximately 10% recombination. Smeltzer found one of the two weak genes (U or V), which he identified in Minturki, also associated with the same linkage group. Crosses between Minturki and other varieties possessing postulated weak genes, however, were not available in his or the current investigations and the relationship between the genes in Minturki and other weak genes could not be established. Presumably U or V may be identical with X. Verification of the postulated genotype for Turkey 10097, where two weak genes are invoked, would necessitate initially isolation of the two separate homozygous genotypes carrying resistance genes in F, lines exhibiting the appropriate levels of infection from a cross with a susceptible variety such as Baart. Final proof would depend on the isolation of homozygous highly resistant or immune F’, lines due to gene interaction when these two genotypes were crossed. It might also reasonably be expected that isolated weak genes similar to those proposed in these studies might be found alone in certain instances. Tisdale et al. (1925) in investigations on varietal resistance in hard red spring, hard red winter, soft red winter and white classes of wheat found that certain varieties and strains in all these classes consistently showed between 10 and 30% infected heads in a series of tests at different stations over a period of two years or longer. The percentage of bunted plants would undoubtedly be higher than this, but there are definite indications that genes permitting only a moderate number of infected plants were responsible for the observed varietal reactions. Likewise El Khishen and Briggs (1945) reported that unpublished data of Mackie and Briggs in 1920 showed that six out of 956 varieties and strains of wheat tested had a bunt infection of between 20 and 25° indicating the presence of a weak gene similar in effect to the X factor. Similarly varieties showing around 40 to 50% infection would indicate the presence of genes similar in effect to the W or Y factors. The group of Turkey wheat selections from the same source as those studied in the present investigations possess weak genes either entirely or in combination with strong ones as they have been discovered in Turkey 10015, 10016, 10095 and 10097. The presence of weak genes in combination with a strong one (R) has been postulated in Turkey 10095 and this parallels the presence of weak genes in combination with the R and T genes in Turkey 10016. Previous investigations on Turkey types and strains of wheat have revealed that, in spite of general morphological similarities, they differ considerably in genotype for bunt resistance. Sherman and Cooperatorka have been found BE. P. BAKER 207 to possess the Martin gene, Turkey 1558, 1558B, 2578, 3055 and Oro the T gene, Rio the R gene, Turkey 10015 the X and Y genes, Turkey 10016 the R, T, X and Y genes and Minturki the U and V genes. Therefore all except the H gene have been found in wheats of this type. Hence the unique genotypes reported in Turkey 10095 and 10097 in the present instance are not unexpected. Clark and Bayles (1935) reported that Turkey is the name applied to a rather morphologically similar group of wheats introduced at different times from widely scattered and isolated areas of the Crimea region in Southern Russia into the United States. Morphologically distinct strains given varietal nomen- clature have been isolated from these wheats and all strains of this variety have been noted as lacking uniformity. Evans and Janssen (1922) reported that South Dakota 144 wheat from which the selections currently studied were made followed the general historical pattern of introduction from the northern areas of the Crimea, whence it was introduced into Kansas about 1875, and was subsequently given the accession number which it bears by the South Dakota Agricultural Experiment Station. In tests reported by Kiesselbach and Anderson (1930) significant differences in cold survival and minor differences in heading and ripening dates were observed among the selections of South Dakota 144, indicating physiological differences. Since the genotypes of Turkey 10015, 10095 and 10097 were different at Davis, California using a single race of bunt, it might reasonably be expected that they would not behave in an identical manner to different bunt races. The only tests on the differential behaviour of these varieties are those reported by Kiesselbach and Anderson. To race VI of 7. foetida, Turkey 10015 showed 8-9% infection whilst the infections in Turkey 10095 and 10097 were higher, being 25-0 and 21-7% respectively. To race X (7. caries) both Turkey 10095 and 10097 were immune but Turkey 10015 showed 7-3% infection. There was little differential behaviour between Turkey 10095 and 10097 and in only two instances were minor differences reported. To one collection of 7. foetida 10095 showed 4-7°% infection whilst 10097 was immune; in the case of one collection of T. caries the respective reactions were immunity and 4-2 °% infection. Tests against several races of the pathogens with varieties known to possess the same gene for resistance to one race will frequently reveal the presence of additional resistance factors. On this basis Holton and Briggs (1950) presented evidence indicating that Martin possessed at least two genes for resistance to a combination of physiologic races of T. caries and T. foetida. Schaller, Holton and Kendrick (1960) showed that a second gene (M,) in Martin was inherited independently of M and was also different from M in being incompletely dominant in inheritance. The varieties Baart and Federation, susceptible to race T-1, apparently possessed the M, gene, since their race reactions were identical to that of the isolated M, gene from Martin. The type of gene interaction whereby a combination of two weak genes are postulated to confer high resistance or immunity to a pathogen is of interest. The interaction may be an additive effect or may involve complementary gene action. The crown rust resistance of the oat variety Bond has been found to be based on complementary dominant gene interaction (Baker and Upadhyaya, 1966 (in press)). From Bond the two factors concerned have been isolated in separate lines (Baker, 1966 (in press)). The phenotypes shown by wheat plants when seed inoculated with the bunt organisms is obviously the result of a complex interaction between genotype and environment. Invariably some genotypically susceptible plants do not manifest bunt symptoms. On the other hand Briggs (1929) indicated that genetically resistant plants, as indicated by progeny tests, occasionally may become infected. He demonstrated also the presence of minor genes which modify bunt resistance. Little is known of the mechanism in the host con- ferring resistance in bunt. A pertinent observation is that if the maturity of the resistant variety Florence is delayed by cutting back the initially formed 908 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS spikes, occasional late maturing tillers show bunted grains (Waterhouse, personal communication). This suggests that a differential growth rate between the host and pathogen after infection confers resistance. In this connection there is also lack of knowledge on the significance of the fact that all tillers in a susceptible plant frequently do not show bunt symptoms. Until basic information is obtained on the nature of bunt resistance in wheat it is impossible to specify the phenomena whereby a homozygous genotype may, in the case of weak genes, permit a moderate percentage of plants to fail to manifest bunt symptoms or those whereby a heterozygous genotype in the case of certain strong genes, such as the R factor, permits approximately only half the plants to show symptoms at maturity. Genotype and environment interactions might reasonably be expected to be more variable in the case of weak genes. In this connection Smeltzer (1952) found that Minturki (possessing the weak genes U and V) in one season showed an average infection of 20-7%, with a range of from 3-8 to 38-9% in individual rows, the previous highest annual average infection over several seasons being 3°2%. He suggested that the physiology of the plants was disturbed by severe winter conditions in the season showing high average infection such that there was a breakdown in the resistance mechanism in Minturki. Environmental effects on host-pathogen relationships have been reviewed by Tapke (1948) in the case of smuts. Five genes (H, M, X, R and T) for resistance to the one race of bunt have been placed by linkage studies on the same chromosome. It may be argued that the R and T genes which are similar in effect and mode of inheritance represent the same gene which differs in its chromosomal site due to a small inversion so that the two genes are no longer at the same locus on the chromosome. However, Sutherland and Jodon (1934) obtained an average of 4% of bunt in 1930-32 with Rio and 60% with Turkey C.I. 1558. At the same time in the absence of more critical evidence this could be accounted for by the presence of a second gene in Rio. The other three genes (H, M and X) clearly differ from one another and from the R and T genes from their reaction to a single race with regard to the level of infection they permit, their mode of inheritance with regard to dominance or the fact that they exhibit a differential reaction to certain races. Sears, Schaller and Briggs (1960) by monosomic analysis placed the M gene on chromosome XIII (2A). Hence the five linked genes of the seven definitely identified conditioning resistance to race T-1 are located on this chromosome belonging to the A genome in hexaploid wheats and contributed by a prototype of the present-day diploid wheat species, Triticum aegilopoides (Link) Bal. This evidence suggests that diploid Triticum species may possess previously unrecorded genes for bunt resistance capable of being transferred by amphidiploidy to hexaploid wheat. Several investigators have reported that T. monococcum L. (2n=14) is generally highly resistant to bunt, but a few instances are noted where it was quite susceptible. It is also of interest that T. timopheevi, Zhuk. containing the A and a modified B genomes, was immune to all bunt races to which it was tested by Rodenhiser and Holton (1945). It is possible that weak genes which have been identified in Turkey wheats on the basis of their segregation pattern to race T-1 may confer immunity to certain races of bunt in the Crimean area. It is also feasible that they may confer immunity to new races as they arise in North America and elsewhere. Hence, although somewhat more difficult to manipulate in a breeding programme on their behaviour to race T-1, they may well have a place in breeding for resistance to particular races of the pathogens. In this connection combinations of single genes may afford protection against numerous physiologic races identified on the accepted differential varieties. Briggs and Holton (1950) concluded that the Martin and either the Turkey or Rio genes together gave E. P. BAKER 209 protection against all 25 then known races. For this reason Selection C.I. 12242, which was assumed to possess both the Rio and Martin genes, was resistant to all races. Schaller and Briggs (1953) reported that two gene: (M and T) in combination afforded protection against 25 out of 31 races then recognized. A combination of the five linked genes (H, M, X, R and T in this order on the chromosome) would provide a potent resistance against all bunt races. The genotype of Turkey 10016 includes the genes T, X and R. By a back- crossing programme involving Turkey 10016 and Martin, using the former variety as the recurrent parent and selecting for resistance to a race such as L-8 or T-16, which is virulent on Turkey 10016, but incapable of attacking varieties possessing the M gene (Briggs and Holton, 1950), the genes M, X, R and T could be combined. Ifa race capable of attacking this combination was detected but to which Hussar was resistant due to the H gene which it possesses, the latter gene could then be incorporated similarly by backecrossing. At the present time there is apparently no bunt race to which Hussar owes its resistance to the combined effect of the M and H genes, and to which Turkey 10016 is susceptible. With the estimated linkage values between these genes it should be possible by crossing Hussar and Turkey 10016 to combine at least the R or T genes with either the M or H genes by selection in segregating generations which are tested with appropriate races. The predicted amount of crossing over would permit their combination in a variety with little difficulty. Once the HMXRT linkage group was established in a homozygous dominant condition in a well-adapted variety the backcross technique could be used in a practical breeding programme. Since monosomic series are available in many agronomic varieties currently cultivated commercially, cytogenetical techniques could also be employed to substitute the chromosome with all the linked genes into these varieties. Acknowledgements Grateful acknowledgement is made to Dr. F. N. Briggs who provided the materials and under whose direction these studies were made. Part of these investigations was carried out during the tenure of a Thomas Lawrance Pawlett Scholarship. References Baxer, H. P., 1966.—Isolation of complementary genes conditioning crown rust resistance in the oat variety Bond. Huphytica, (in press). Baker, E. P., and Upapuyaya, Y. M., 1966.—Studies on the inheritance of rust resistance in oats. IV. Expression of allelomorphism and gene interaction for crown rust resistance in crosses involving the oat variety Bond. Proc. Linn. Soc. N.S.W., (in press). Briaes, F. N., 1926.—Inheritance of resistance to bunt, T%lletia tritica (Bjerk.) Winter, in wheat. J. agr. Res., 32: 973-990. , 1929.—F actors which modify resistance of wheat to bunt, Tlletia triticsx. Hilgardia, 4: 175-181. , 1930a.—Inheritance of the second factor for resistance to bunt, V%lletia tritici, in Hussar wheat. J. agr. Res., 40: 225-232. , 1930b.—Inheritance of resistance to bunt, T%lletia tritict, in White Odessa wheat. J. agr. Res., 40: 353-359. , 1931.—Inheritance of resistance to bunt, Tvlletia triticc, in hybrids of White Federation and Banner Berkeley wheats. J. agr. Res., 42: 307-313. , 1932a.—Inheritance of resistance to bunt, Tlletia tritici, in hybrids of White Federation and Odessa wheat. J. agr. Res., 45: 501—505. . 19326.—Inheritance of resistance to bunt, J7lletia tritici, in crosses of White Federation with Turkey wheat. J. agr. Res., 44: 131-136. , 1934.—Inheritance of resistance to bunt, Tlletza tritzct, in Sherman and Oro wheat hybrids. Genetics, 19: 73-82. , 1936.—Inheritance to resistance to bunt, Tilletia tritici, in hybrids of Turkey wheats, C.I. 1558B and 2578. Hilgardia, 10: 19-25. , 1940.—Linkage between the Martin and Turkey factors for resistance to bunt, Tilletia tritici. J. Amer. Soc. Agron., 32: 539-541. Brices, F. N., and Hotton, C. 8., 1950.—Reaction of wheat varieties with known genes for resistance to races of bunt, Tilletia caries and T. foetida. J. Amer. Soc. Agron., 42: 483-486. 210 INHERITANCE OF RESISTANCE TO BUNT IN TURKEY WHEAT SELECTIONS CuarkK, J. A., and Bayvtss, B. B., 1935.—Classification of wheat varieties grown in the United States. U.S. Dept. Agric. Tech. Bull. 459. Ex KutsHen, A. A., and Briaes, F. N., 1945.—Inheritance of resistance to bunt (T%lletia caries) in hybrids with Turkey wheat selections C.I. 10015 and 10016. J. agr. Res., 71: 403-413. Evans, A. T., and JANSSEN, G., 1922.—Winter wheat in South Dakota. S. Dak. agric. Expt. Stat. Bull. 200. KiessELBAcH, T. A., and ANDERSON, A., 1930.—Breeding winter wheats for resistance to stinking smut. Nebr. agr. Expt. Stat. Res. Bull. 51. Kureer, J., 1965.—Failure of hexachlorobenzene to control common bunt of wheat. Nature, 206: 1219-1220. RopENHISER, H. A., and Hotton, C. §., 1937.—Physiologic races of Tilletia tritict and T. levis. J. agr. Res., 55: 483-496. 1945.—Distribution of races of Tilletia caries and Tilletia foetida and their relative virulence on certain varieties and selections of wheat. Phytopath., 35: 955-969. SCHALLER, C. W., and Briaes, F. N., 1955.—Linkage relationships of the Martin, Hussar, Turkey and Rio genes for bunt resistance in wheat. J. Amer. Soc. Agron., 47: 181-186. ScHaLLerR, C. W., Hotton, C. S., and Kenpricx, E. L., 1960.—Inheritance of the second factor for resistance to bunt, Tilletia caries and T. foetida, in the wheat variety Martin. J. Amer. Soc. Agron., 52: 280-282. Sears, FE. R., ScoHaLLteR, C. W., and Briaes, F. N., 1960.—Identification of the chromosome carrying the Martin gene for resistance of wheat to bunt. Canad. J. Genet. Cytol., 2: 262-267. SMELTZER, D. G., 1952.—Inheritance of resistance to race T—1 of Tilletia caries in Minturki and Cooperatorka wheats. J. Amer. Soc. Agron., 44: 529-533. STANFORD, E. H., 1941.—A new factor for resistance to bunt, Vulletia tritici, linked with the Martin and Turkey factors. J. Amer. Soc. Agron., 33: 559-568. SUTHERLAND, J. L., and Jopon, N. E., 1934.— Resistance of wheat varieties at Moccasin, Montana and North Platte, Nebraska. J. Amer. Soc. Agron., 26: 296-306. TapKs, V. F., 1948.—Environment and the cereal smuts. Bot. Rev., 14: 349-412. TIsDALE, W. H., Martin, J. H., Briaes, F. N., Mackir, W. W., Wootman, H. M., STEPHENS, D. E., Gatnes, E. F., and Stevenson, F. J., 1925.—Relative resistance of wheat to bunt in the Pacific coast states. U.S. Dept. Agric. Bull., 1299. RHINONYSSINE NASAL MITE INFESTATIONS IN BIRDS AT MITCHELL RIVER MISSION DURING THE WET AND DRY SEASONS ROBERT DOMROW Queensland Institute of Medical Research, Brisbane [Read 30th November, 1966] Synopsis The tropical Mitchell River Mission has a sharply defined wet and dry season each year, almost 90% of the annual average rainfall of 48 inches falling between December and March. Collections of birds during each season showed no obvious difference either in terms of absolute numbers or species composition, except for a slightly broader spectrum in the “ dry ” (59% of the species were seen in both seasons, 25% only in the “dry” and 16% only in the “ wet’). Any disparity is explainable by (i) natural factors such as seasonal migration and nomadism ; and (11) sampling bias: the country is largely impassable in the “ wet’, and, paradoxically, it is therefore easier both to collect water-birds in the “ dry ” and perching birds in the “ wet ”’. This constancy is paralleled by the rhinonyssine mite populations in the nasal passages of these birds. The same mite genera and species are present in both seasons, and infestation rates and numbers of mites per infested bird are constant. Of 108 species of birds examined, 43 had nasal mites; the proportion in the “ wet” was 28/77 and in the “ dry ” 29/82. Of 841 individual birds examined, 129 had nasal mites; the proportion in the “ wet ”’ was 65/476 and in the “ dry ” 64/365. The average number of mites in an infested bird was five in the “ wet ” and four in the “dry”. These overall statistics could reflect the relatively constant micro- climate in the warm, moist nasal passages compared with the extremes outside, where noon shade temperatures over 100°F and relative humidities below 20% are common in the “ dry ”’. A list of the 128 species of birds recorded at the Mission is appended, showing the species recorded, or likely to be recorded as harbouring rhinonyssine nasal mites. INTRODUCTION The belief of Victorian virologists that Murray Valley encephalitis had infiltrated from north Queensland stimulated a survey of human sera for neutralizing antibodies to MVE. This showed their highest incidence around the Gulf of Carpentaria, and the Mitchell River Mission was selected for intensive study. The virus has since been isolated from mosquitoes caught there, and periodical visits have confirmed the significance of the area, where a field station has recently been opened (20th Annual Report, Queensland Institute of Medical Research, 1965). In an expansion of this programme, several species of birds have already been shown, by direct isolation of viruses, to act as reservoirs of infection. The study of blood-sucking arthropods other than mosquitoes as possible natural vectors is also contemplated, but this will be hampered for some time by the infant state of systematics in almost all groups concerned. As a beginning, the intranasal mites of birds are receiving attention. The present note, a comparison of the bird faunas and their intranasal mite populations in the ‘‘ wet ’’ and the ‘‘ dry ”’, is based on 10 weeks intensive field collecting, divided almost equally between the dry seasons (October- November) of 1964 and 1965, and the wet seasons (March-April) of 1965 and 1966. Results to date have been most gratifying, and a wealth of new records and Species is under study (Domrow, 1966, and in ms.). PROCEEDINGS OF THE LINNEAN SocteTy or NEw SoutrH Watss, Vol. 91, Part 3 212 RHINONYSSINE NASAL MITE INFESTATIONS THE LOCALITY To summarize Standfast (1965), the Mitchell River Mission cattle station (15°28’S, 141°40’E) is centred 16 miles from the west coast of Cape York Peninsula in a very flat, low-lying area of tropical tussock grassland and tropical woodland. It is typified by scrubby eucalypts and many coarse grasses, with fresh-water mangrove (Barringtonia gracilis)* and cabbage-tree palms (Livistona australis) common along the water-courses. The annual rainfall averages 48 inches, of which 42 inches fall between December and March. During the ‘¢ wet ’’, the rivers and creeks flood and large areas of low-lying country become fresh-water swamps, which, overgrown with the ubiquitous and very dense and tall grasses, provide a breeding ground for a wide variety of water-birds. During the “‘ dry ’’, this grass largely dies (or is burnt) away, leaving much bare earth, while the creeks are restricted to a series of water-holes, and the swamps recede to form small lagoons or dry out completely. These residual pools, with their meagre fringe of greenery, become the foci of the activities of large numbers of birds, which come to water throughout the day—this is not surprising as frequently the shade temperature exceeds 100°F and the relative humidity falls below 20°, at noon. THE BIRDS Tables 1 and 2 are confined to the birds I have actually examined for nasal mites, and provide a comparison between the ‘‘ wet’ and the “dry”. In both, the host data are broken down to ordinal level. Table 1 is concerned with the numbers of species of birds examined and found infested. Table 2 details the total numbers of birds examined, the numbers found infested and the numbers of mites per infested bird. TABLE | Numbers of species of birds examined and found infested with Rhinonyssinae at Mitchell River Mission Species examined Species infested Order “Wet” “Dry” Both Total “Wet” “Dry” Both Total 2 6 3 bo Columbiformes . . Ke 4 1 4 Gruiformes Podicipiformes . . Charadriiformes Ciconiformes Anseriformes Pelecaniformes . . Faleconiformes . . Strigiformes Psittaciformes .. Caprimulgiformes Coraciiformes Cuculiformes Passeriformes .. oo = KOO PE OeNWNON (SY) PNT TE WNADOR eH > dob oe > (JU) ND > WADE AN MHwWH H bo bo ome bo in Total -~] «J GO bo oO — — (=) ee) bo (o/2) bo Neo) — is nS (SU) It is evident, both in terms of species composition and absolute numbers of individual birds, that the “‘ wet’ fauna parallels the ‘“‘ dry ’’ very closely (there are several large and permanent water-holes in the area), but with a slightly broader spectrum in the ‘‘dry”’. This is confirmed by consulting the complete list below of the birds now known to occur at Mitchell River: 75 (59°) of the 128 species were listed in both seasons, with 32 (25°%) noted only in the ‘“‘ dry ’ against 21 (16%) only in the “ wet’. However, although it is impossible * Given in error as Nauclea orientalis by Standfast. ROBERT DOMROW 213 to give a clear-cut explanation in every case—the predatory and water-bound faunas, for example, while nomadic, are undoubtedly more uniform than indicated—there are two factors which would tend to emphasize this slight difference. The first comprises real, or natural differences of two main types. The dry season component includes a group of migratory species (Myristicivora, Mesoscolopax, Hrolia, Stiltia, Scythrops and Myiagra), which, travelling south in September-October and returning north in March-April, could also be expected at the Mitchell River Mission in the “‘ wet’. However, other members of this group (Halcyon sancta, Cacomantis variolosus and Chalcites) have been noted only in the ‘“ wet’’, and Hudynamys and Hdoliisoma in both seasons. Also, many (particularly a large group of nectar-feeders, e.g., lorikeets and honey-eaters) are nomadic, being dependent on flowering plants etc.: Barring- tonia and eucalypts were both flowering profusely in the ‘“ dry’ of 1965. The mistletoe bird, Dicaeum, is another example: this small bird ‘‘ undertakes extensive seasonal movements coincident with the fruiting of the [mistletoe] plants. This may be in spring, summer, autumn, or winter, in different areas ”’ (Keast, 1961). The second is an unreal difference due to collecting bias. This is the direct result of climatic conditions. In the ‘‘ wet’, the roads are closed even to light traffic, and the swamps are overgrown with dense grass and reeds in excess of eight feet tall. Tables 1 and 2 bear out the paradoxes that it is easier both to obtain water-birds in the “‘ dry” (95 of 25 species v. 55 of 18 species) and perching birds in the ‘“ wet ”’ (421 of 59 species v. 270 of 57 species). Further, it is impossible to reach the sea by overland in the ‘‘ wet ”’, so Larus, Haematopus and Haliastur indus have been noted only in the “ dry ”’. THEIR NASAL MITE PARASITES From Tables 1 and 2, the infestation rates (Rhinonyssinae only) are also remarkably similar. Omitting host species examined only in the dry season, and the Cuculiformes, of which only inconclusive numbers were examined, the types of birds infested, the numbers infested and the numbers of mites per infested host remain virtually unchanged from the “‘ wet” to the “dry ”’. TABLE 2 A seasonal comparison of infestation rates by Rhinonyssinae in birds at Mitchell River Mission Number examined Number infested Mites/infested bird Order Ne Dray PS et Dig 7 CONE E Die 9 Columbiformes Be! i 32 32 5 4 3(3) 2(2) Gruiformes .. Py ia 7 Podicipiformes NS ae 2 1 1(1) Charadriiformes .. ws 20 41 5 9 4(3) 6(6) Ciconiiformes Ah ie 19 13 2 16(15) Anseriformes. . i Fi 15 28 1 2(2) Pelecaniformes ae wy 1 4 Falconiformes wi Us 24 8 5 2 6(4) 4(1) Strigiformes .. ne die il 1 Psittaciformes ae in 62 61 4 6 4(—) 4(4) Caprimulgiformes .. ed 1 2 Coraciiformes He aa 44 18 15 2 4(3) 10(—) Cuculiformes. . . - 17 4 2 1 20(-) 1(1) Passeriformes ae Ae 240 144 29 36 4(4) 3(2) Total .. Ge i 476 365 65 64 5(3) 4(3) In the final column, unbracketed figures are derived from the total number of mites present ; bracketed figures apply only to sexually mature mites—the adjustment is not great. Dashes indicate that figures are unavailable, certain common mites having been deep-frozen in the field for subsequent inoculation studies. The bracketed figures for Passeriformes are, for this reason calculated from only 26 birds in the “ wet”? and 31 in the “dry”. 214 RHINONYSSINE NASAL MITE INFESTATIONS The constancy in seasonal occurrence of birds extends to the genera and species of their mite parasites, which are now treated under an ordinal arrange- ment of those hosts known to occur in both seasons. Host-specificity is marked in the Rhinonyssinae. Double rhinonyssine infestations, while not unknown elsewhere in Queensland (Domrow, unpublished data), have not been noted at Mitchell River, although occasional mixed infestations—Rhinonyssinae (Mesostigmata) and Speleogna- thinae (Prostigmata)—are on record. However, speleognathines tend to inhabit portions of the nasal cavities different from those preferred by rhinonyssines. COLUMBIFORMES: The two species of mites recorded (Mesonyssus geopeliae and M. ocyphabus) are members of a widespread and characteristic group of Mesonyssus species peculiar to pigeons (Fain, 1962; Wilson, 1964; Domrow, 1965). CHARADRUFORMES: Both genera and species of mites noted (Rhinonyssus himantopus and Larinyssus benoitt) are widespread parasites of waders (Strandt- mann, 1959; Domrow, 1966). FALCONIFORMES: The one species collected, Ptilonyssus cerchneis, also occurs in falcons in Africa, U.S.A. and U.S.S.R. (Fain, 1957; Strandtmann, 1962; Bregetova, 1964). PSITTACIFORMES : All three species of mites recorded (Mesonyssus trichoglosst, M. kakatuae and M. aprosmicti) are members of a group of Mesonyssus species restricted to this order of birds (Wilson, 1964; Domrow, 1964). CORACHUFORMES: The two species of Mesonyssus (M. daceloae and M. haleyonus) recorded from Alcedinidae are close relatives of the species known from African members of this family (Fain, 1957, 1960; Domrow, 1965). Merops, a bee-eater (Meropidae), is parasitized by a species of Ptilonyssus, P. triscutatus, previously known from this host genus only in Europe and Africa (Fain, 1957). CUCULIFORMES: The species of Sternostoma, S. cuculorum, taken from cuckoos at Mitchell River was originally recorded from the same host family in Africa (Fain, 1957). PASSERIFORMES: Apart from Sternostoma thienponti and Sternostoma sp. from Cracticus and Neositta respectively, and Ruandanyssus sp. from Artamus, the remaining 13 species from 17 birds belong to Ptilonyssus (or the closely related Passeronyssus). These will not be listed in detail, but where overseas hosts have been recorded for these species, they have been closely related indeed to the Australian ones. DISCUSSION One explanation for this remarkable constancy of parasite populations throughout the year would seem to lie in the clemency of the warm, moist microclimate afforded by the nasal passages of the host, compared with the extremes outside. On the other hand, with small arthropods bound absolutely to water, ¢.g., mosquitoes, the wet season fauna differs radically from the dry, when standing waters stagnate and specialized habitats such as temporary pools, tree-holes and leaf axils dry out. From Table 2 it was seen that the incidence of immature stages in both the ‘“‘ wet” and the “dry ”’ was low, necessitating only minor adjustment to the overall numbers of mites per infested host to express their presence. Both sets of figures are given, however, to stimulate enquiry into the question whether the mite populations fluctuate at other times of the year. Further, the tenacity with which these delicate, weakly sclerotized mites keep to their specific micro- habitat (see Addendum), coupled with their low numbers at the periods studied, presents the possibility that the restricted collecting periods may have tapped only maintenance populations, and not have coincided with reproduction peaks. ROBERT DOMROW 215 A List oF MITCHELL RIVER BIRDS As both the wet (W) and dry (D) seasons are considered above, my own data have been slightly augmented by the incorporation of Standfast’s preliminary list (1965) of the dry season fauna, on which I have the following comments. It is doubtful if more than one species of crow (Corvus cecilae) occurs at the Mission : eye-colour varies tremendously from white, through parti-coloured. (blue and white), to dense blue-black, while the down of all specimens is uniformly white. With this one deletion (C. bennett), I have had a further 41 of Mr. Standfast’s species in the hand and would accept them, with two reservations. Firstly, the smaller of the two species of cuckoo-shrikes occurring at the Mission admittedly appears white-breasted in the field, but in the hand it is distinctly grey-tinted and identifiable as Coracina papuensis rather than C. hypoleuca. Secondly, the birds pointed out to me by Mr. Standfast as ‘* banded plovers ” proved in the hand to be Australian pratincoles (Stiltia isabella), not Zonifer tricolor. Of the remaining 15 species, I can confirm seven by sighting, track or call (Dromaius, Burhinus, Xenorhynchus, Pelecanus, Haliastur indus, Falco cenchroides and Scythrops), but have not noted the others (Alectura, Phaps, Porphyrio, Dupetor, Alcyone, Cyrtostomus, Aegintha and Ptilonorhynchus). I can now add another three orders (Podicipiformes, Strigiformes and Caprimulgiformes) and 72 species to this list. Of these, five (Synoicus, Larus, Haematopus, Himantopus and Notophoyx pacifica) are based on adequate sightings in the field and the remainder on specimens in the hand. These latter, in cases of doubt, were sent to the Queensland Museum, where Mr. D. P. Vernon kindly checked them against reference skins. I do not think many species, at least of land birds, remain undetected in the area, but I have glimpsed snipe, Swifts, swallows and possibly a fantail. The following arrangement is that of the 9th edition [1958] of Leach’s An Australian Bird Book. The rhinonyssine mite species printed in italics have been recorded at Mitchell River, while those in Roman type may be expected to occur there, having been recorded from the indicated host elsewhere in Australia (or New Guinea in the case of Anas superciliosa, see Wilson, 1964).* CASUARIIFORMES Dromaius novaehollandiae WD GALLIFORMES Alectura lathami D Synoicus australis WD COLUMBIFORMES Myristicwora spilorrhoa D Mesonyssus sp. “ A ”’ Mesonyssus sp. “ B” Geopelia placida WD Mesonyssus geopeliae Geopelia cuneata WD Geopelia humeralis WD Mesonyssus geopeliae Phaps chalcoptera D Mesonyssus phabus Geophaps scripta W Mesonyssus ocyphabus Ocyphaps lophotes W Mesonyssus ocyphabus Mesonyssus columbae GRUIFORMES Porphyrio melanotus D Grus rubicundus WD * Since writing this, I again visited Mitchell River for three weeks in October-November 1966, adding eight more species to the list, making 136 in all. These additions are indicated by ““d” in the central column. 216 RHINONYSSINE NASAL MITE INFESTATIONS PODICIPIFORMES Podiceps ruficollis D Rhinonyssus poliocephali CHARADRIIFORMES Chlidonias hybrida WD Sterna bergw d Larinyssus orbicularis Larus novaehollandiae Haematopus ostralequs Hrythrogonys cinctus Lobibyx miles Charadrius alexandrinus Charadrius melanops Himantopus leucocephalus Recurvirosira novaehollandiae Numenius madagascariensis Mesoscolopax minutus Inmosa limosa Hrolia ruficollis Hrolia acuminata TIrediparra gallinacea Stiltia isabella Burhinus magnirostris Hupodotis australis CICONIIFORMES Threskiornis molucca WD Threskiornis spinicollis WD Plegadis falcinellus WD Platalea regia WwW Platalea flavipes Ww Xenorhynchus asiaticus D Eigretta intermedia Wd Eigretta alba WD Notophoyx novaehollandiae WD Mesonyssus belopolski Notophoyx pacifica Wd Notophoyx picata WD Mesonyssus belopolskiw Nycticorax caledonicus WD Dupetor flavicollis D ANSERIFORMES Anseranas semipalmata WD Nettapus pulchellus WD Dendrocygna arcuata D Rhinonyssus rhinolethrus Tadorna radjah WD Anas superciliosa D Rhinonyssus rhinolethrus Anas gibberifrons D PELECANIFORMES Phalacrocorax sulcirostris WD Phalacrocorax melanoleucus WD Anhinga novaehollandiae WD Pelecanus conspicillatus WD FALCONIFORMES Accipiter fasciatus Wd Aquila audax WD Haliaeetus leucogaster WD Haliastur sphenurus WD Haliastur indus D Milous migrans WD Falco berigora WD Ptilonyssus cerchneis Falco cenchroides WD Ptilonyssus cerchneis STRIGIFORMES Ninox novaeseelandiae WwW Rhinoecius cooremani Ninox connivens D SecSvedteanus eguuy Rhinonyssus himantopus Rhinonyssus himantopus Rhinonyssus coniventris Rhinonyssus minutus Rhinonyssus himantopus Rhinonyssus himantopus Rhinonyssus sp. Larinyssus benoiti ROBERT DOMROW 217 PSITTACIFORMES Trichoglossus moluccanus WD Mesonyssus trichoglossi Psitieuteles versicolor D Calyptorhynchus banks WD Mesonyssus kakatuae Kakatoe galerita WD Kakatoe roseicapilla WD Mesonyssus kakatuae Kakatoe sanguinea D Aprosmictus erythropterus WD Mesonyssus aprosmicti CAPRIMULGIFORMES Podargus strigoides WD CoRACIIFORMES Hurystomus orientalis WD Alcyone azurea D Dacelo leachi WD Mesonyssus daceloae Halcyon pyrrhopygia WD Mesonyssus halcyonus Halcyon sancta Ww Mesonyssus halcyonus Halcyon macleayt WD Mesonyssus halcyonus Merops ornatus WD Ptilonyssus triscutatus Sternostoma cooremani CUCULIFORMES Cuculus saturatus W Cacomantis pyrrhophanus WD Sternostoma cuculorum Cacomantis variolosus Ww Sternostoma cuculorum Chalcites plagosus Wd Scythrops novaehollandiae D Hudynamys orientalis WD Centropus phasianinus WD PASSERIFORMES Hirundo neoxena d Cas angrensis Ptilonyssus echinatus Microeca fascinans -WD Piilonyssus microecae Microeca flavigaster WD Ptilonyssus microecae Rhipidura leucophrys WD Ptilonyssus maccluret Sternostoma laniorum Myiagra rubecula D Sternostoma laniorum Ruandanyssus terpsiphonei Seisura inquieta D Ptilonyssus terpsiphonei Coracina novaehollandiae WD Ruandanyssus terpsiphonei Coracina papuensis WD Edoliisoma tenutrostre WD Lalage tricolor WD Ptilonyssus cractici Ruandanyssus terpsuphoner Smicrornis flavescens Wd Gerygone palpebrosa WD Ptilonyssus sp. Cisticola exilis WD Malurus melanocephalus WD Ptilonyssus maluri Artamus cinereus WD Ruandanyssus sp. Grallina cyanoleuca WD Ptilonyssus grallinae Pachycephala rufiventris WD Ptilonyssus motacillae Ruandanyssus terpsiphonei Gymnorhina tibicen Wd Ptilonyssus cractici Sternostoma thienponti Cracticus nigrogularis WD Sternostoma thienponti Cracticus quoyt WD Sternostoma thienponti Neositta striata WwW Sternostoma sp. Climacteris melanota d Dicaeum hirundinaceum D Ptilonyssus dicael Pardalotus rubricatus WwW Cyrtostomus frenatus D Ptilonyssus cinnyris Melithreptus albogularis WwW Ptilonyssus meliphagae Myzomela pectoralis WD Ptilonyssus gliciphilae Gliciphila fasciata D Gliciphila indistincta D Ptilonyssus gliciphilae Conopophila albogularis d Meliphaga fusca WwW Meliphaga flava WD Ptilonyssus stomioperae 218 RHINONYSSINE NASAL MITE INFESTATIONS Stomiopera unicolor WD Piilonyssus stomioperae Entomyzon cyanotis WD Ptilonyssus philemoni Philemon citreogularis D Pitlonyssus philemona Mirafra javanica WD Ptilonyssus capitatus Steganopleura bichenovir WwD Donacola castaneothorax WD Aegintha temporalis D Bathilda ruficauda W Poephila atropygialis WD Poephila personata WD Poephila gouldiae WwW Sternostoma tracheacolum Oriolus flavocinctus WD Ptilonyssus trouessarti Sphecotheres flaviventris WD Ptilonyssus sphecotheris Chibia bracteata WD Passeronyssus dicrurt Ptilonorhynchus violaceus D Chlamydera nuchalis WD Ptiilonyssus sphecotheris Corvus cecilae WD Acknowledgements Our Institute is most grateful to the Anglican Mission at Mitchell River for many courtesies. The Government Botanist, Mr. 8S. L. Everist, kindly identified the Barringtonia from a kodachrome: its foot-long, pendulous, red inflorescences are conspicuous in the ‘dry ”’. References Brecetova, N. G., 1964.—Some problems of evolution of the rhinonyssid mites. Report presented at the First International Congress of Parasitology. Nauka, 1-7. Domrow, R., 1964.—The genus Mesonyssoides in Australia (Acarina: Laelapidae). J. ent. Soc. Qd., 3: 23-29. , 1965.—New laelapid nasal mites from Austrahan birds. Acarologia, 7: 430-460. , 1966.—Some mite parasites of Australian birds. Proc. Linn. Soc. N.S.W., 90: 190-217. Fatn, A., 1957.—Les acariens des familles Epidermoptidae et Rhinonyssidae parasites des fosses nasales d’oiseaux au Ruanda-Urundi et au Congo belge. Ann. Mus. roy. Congo belge, 60: 1-176. , 1960.—Acariens nasicoles récoltés par le Dr. F. Zumpt en Rhodésie du Nord et au Transvaal. Description de trois espéces nouvelles. Rev. Zool. Bot. afr., 62: 91-102. , 1962.—Les rhinonyssides parasites des pigeons (Acarina: Mesostigmata). Rev. Zool. Bot. afr., 65: 305-324. Kzast, A., 1961.—Bird speciation on the Australian continent. Bull. Mus. comp. Zool. Harv., 123: 305-495. StanprFast, H. A., 1965.—Notes on the birds of Mitchell River. Qd. Nat., 17: 91-94. STRANDTMANN, R. W., 1959.—New records for Rhinonyssus himantopus and notes on other species of the genus. J. Kansas ent. Soc., 32: 133-136. , 1962.—A ptilonyssid mite from the sparrow hawk, Falco sparverius (Acarina : Rhinonyssidae). Proc. ent. Soc. Wash., 64: 100-102. Witson, N., 1964.—New records and descriptions of Rhinonyssidae, mostly from New Guinea (Acarina: Mesostigmata). Pacific Insects, 6: 357-388. ADDENDUM Since writing the above, it has become desirable to explain the method of collection, and to detail the parasitope of these mites. All the material was taken after the classical method outlined by Fain (1957), the premier student of these mites: ‘‘ La récolte des Acariens a été pratiquée généralement peu de temps apres la mort de ?Oiseau . . . le bee est largement ouvert. Au moyen dune paire de ciseaux a mors fins on découpe le palais, le plus prés possible du bec et sur une grande longueur de fagon a bien exposer la région qui correspond aux narines. Les tissus excisés sont... examinés ultériewrement ... Pour examiner les narines par Vintérieur il est souvent nécessaire de disséquer les cornets situés profondément ou les lamelles cornées qui cachent plus ou moins leur orifice interne . . ROBERT DOMROW 219 Les Rhinonyssidés vivent . . . dans les narines, on peut cependant les rencontrer ausst dans le mucus qui recouvre les cornets, mais ils ne s’aventurent jamais trés loin a Vintérieur des fosses nasales . . . les Rhinonyssidés sont animés de mouve- ments lents et ne se déplacent probablement que trés peu.” The same author (1965, Ann. Parasit. hum. comp., 40: 317-327) continues : ‘““ De toutes les formes Wacariase interne, la plus répandue est celle des voies respiratoires. On la rencontre principalement chez les Mammiferes et les Oiseaux, mais elle existe aussi chez les Serpents, les Batraciens et méme chez certains Invertébrés tels que les Mollusques pulmonés ... Tous ces Acariens des votes respiratoires sont spécifiques, non seulement en ce qui concerne Vhéte, mais encore pour la région souvent trés limitée qwils occupent dans Vappareil respiratoire .. . Les Acariens dela... famille... des Rhinonyssidae sont hématophages comme la plupart des Mésostigmates ectoparasites dont ils dérivent probablement. On les rencontre .. . dans les régions antérieures des fosses nasales, mais uniquement sur les parties les plus vascularisées des cornets. Une espéce cependant (Sternostoma tracheacolum a envahi les bronches et les poumons ou elle se nourrit @ailleurs également de sang.” Any tendency, therefore, based only on clinical veterinary experience with this species (it causes severe respiratory complications in cage birds, see Murray, 1966, Aust. vet. J., 42: 262-264), to believe that rhinonyssine mites in general are endoparasites whose major populations occur in the deeper respiratory organs (tracheae, bronchi, lungs and air sacs), with only the fringe of those populations evident in the nasal passages, should be resisted. Avian nasal mites are just that, and all specialists I have questioned on this point agree, having dissected birds in the field, that rhinonyssines are not found, except aS rarities and S. tracheacolum apart, in the internal respiratory tract (see also Maa and Kuo, 1965, J. med. Hnt., 1: 395-401). By this method it is possible to collect virtually every mite present in the nasal passages of each bird. The population data given above are therefore both quantitative and at least suggestive that the populations of these delicate mites are independent of the two major macroenvironmental factors, relative humidity and temperature. Nor does it seem likely that these factors would affect the mite populations by way of their hosts, since avian organ systems are capable of considerable physiological regulation, even in extreme climatic conditions (Thomson, 1964, ‘‘ A New Dictionary of Birds’, Nelson, London— see entries on drinking, excretion, heat regulation, respiratory system, efc.). To conclude, rhinonyssine mites are essentially the sole occupants of a specialized parasitope affording an unchanging microclimate and a food supply of constant quality. Reasonably stable populations are therefore not inconsistent with this situation. However, I know of no published data to support this, and so let the original text stand. Corrigenda These Proceedings, Vol. 90, Part 2, page 199, line 11 from bottom, for “‘ pale-yellow robin, Eopsaltria capito”’’ read “‘\lemon-breasted flycatcher, Microeca flawigasier”. Page 208, line 9, for ‘‘ Callistemon”’ read “‘ Barringtonia gracilis’’. Also amend host data accordingly in Synopsis and captions to Figs 28 and 29. INTRASPECIFIC POLYPLOIDY IN HYPOPTERYGIUM ROTULATUM (HEDW.) BRID. HELEN P. RAMSAY School of Biological Sciences, University of Sydney (Plates vi—ix) [Read 30th November, 1966] Synopsis Intra specific polyploid races with chromosome numbers of n=9, 18, ca. 27 and 36 were discovered in plants of Hypopterygium rotulatum growing mixed together on the same log at Mt. Wilson, and a population with n=18 was located a mile away. Cytological behaviour at meiosis was studied, and morphological characteristics of the plants were compared. Spore mother cells with normal meiosis were found in all chromosome races, but some higher polyploids showed a high proportion of irregularities. Few morphological differences between the various plant groups were detected. Some increase in the size of spore mother cells and greater variability in size of spores occurred in high polyploids. The leaf cells were more variable and longer in the plants with chromosome numbers of n=18 and 36. Plants with a chromosome number of n=9 were dioecious while the others were monoecious. INTRODUCTION The HYPOPTERYGIACEAE is a well-defined tropical family of mosses, Hypopterygium being the largest genus with about 70 species (Brotherus, 1924). Hypopterygium rotulatum (Hedw.) Brid. in the sub-genus Eu-hypopterygium, section Pseudo-Tamariscina, is a very variable species (Sainsbury, 1955) evident from the large number of specimens described originally as separate species but now moved into synonymy. The upright almost dendroid gametophytes are produced from a creeping secondary protonema, a number of plants arising along its length. Some of the plants are perennial although new gametophytes arise from the protonema, or new shoots form from the old plants each year. The complanate arrangement of the leaves, and the ventrally placed amphi- gastria characterize this family. H. rotulatum has been recorded in the literature as dioecious (Brotherus, loc. cit.) each female gametophyte bearing several capsules, usually only one from each perichaetium. Shimotomai and Koyama (1932) obtained a mitotic count of n=18 from gametophyte tissue in the synoecious species H. japonicum Mitt. belonging to the sub-genus Eu-Hypopterygium, section Aristifolia. No other chromosome numbers have been recorded for this family. MATERIALS AND METHODS The specimens of Hypopterygium rotulatum (Hedw.) Brid. were collected from two areas about one mile apart at Mt. Wilson, N.S.W. at an altitude of about 3,200 ft. in shaded rainforest. The majority were growing on a log which was covered for more than six feet on its sides and underneath by H. rotulatum and Pterygophyllum dentatum. Plants of Hypopterygiwm were growing on rotting logs, earth and rocks on the banks of a stream. As some variation in the sizes of plants was noticed, groups of plants were kept separate. Specimens were PROCEEDINGS OF THE LINNEAN Society oF New SoutH Watss, Vol. 91, Part 3 HELEN P. RAMSAY 22 kept in glass dishes for two to three weeks in the laboratory until meiosis was completed, then plants were pressed. These voucher specimens have been retained in the Ray Herbarium, Botany Department, University of Sydney (for numbers see Table 1). Smear preparations of meiotic stages in spore mother cells were obtained using the method outlined in Ramsay (1964, 1966). Drawings and photomicro- graphs were made using Zeiss microscope equipment and phase contrast microscopy as in Ramsay (1966) and are reproduced here at a magnification of x2,700 for drawings and at quoted magnifications for the photomicrographs. OBSERVATIONS In Table 1 it can be seen that a series of chromosome numbers was obtained for the 12 different groups of plants of Hypopterygium rotulatum examined. The chromosome numbers of n=9, 18 ca. 27 and 36 represent a polyploid series in plants growing in the same locality and in close proximity. The results deal separately with the chromosome numbers and behaviour at meiosis, and with the comparison of taxonomic features of plants from each of the chromosome groups. TABLE 1 Chromosome numbers in Hypopterygium rotulatum Date Voucher* Chrom. . ; Collected Number See Number Locality Figures Plates 16-464 8/64 D 9 Log, waterfall 1 WAI 6—9-65 36/65 D 9 Log, waterfall 2-3 V1.2 16—4—64 8a/64 M 18 Log, waterfall 4, 6 V1.3 15-64 33/64 M 18 Happy Valley 5 VI.4 33a/64 336/64 : 15-64 22/64 M ca. 27 Rocks, waterfall 7-9 == 16—4—64 8d/64 M Cas 2 Log, waterfall VI1.5-11 16—4—64. 8e/64 M 36 Log, waterfall 10-11 Vale =3 16—4-64 86/64 M 36 Log, waterfall = = 16—4_64 8/64 M 36 Log, waterfall — VILI.4 16—464 8f /64 M 36 Log, waterfall — V1II.5-8 16—4—64 16/64 M 36 Rocks, waterfall == —_ M=monoecious. D=dioecious. All specimens were collected from Mt. Wilson (3,200 ft.) in New South Wales. = * Specimens deposited in the Ray Herbarium, Botany Department, University of Sydney. CYTOLOGICAL BEHAVIOUR H. rotulatum (8/64, 36/65) n=9 (Text-figs 1-3. Plate VI, 1-2). Both these dioecious populations were collected from the same log, the former in April, 1964, the latter in September, 1965 (Table 1). A chromosome number of n=9 was observed at Metaphase I of meiosis (Text-figs 1-2. Plate VI, 1-2). Hight of the bivalents were of almost uniform size, but one was smaller and disjoined precociously in the majority of cells (Text-fig. 2). Its Size and precocious behaviour are similar to that found in a number of other mosses (Vaarama, 1953, 1956, Steere, 1954, etc., Yano, 1957, Khanna, 1960, etc., and Ramsay, in a number of Australian mosses unpubl.). At Anaphase I, nine chromosomes were observed to pass regularly to each pole (Text-fig. 3). Tetrad formation was normal and no lag of chromosomes was seen. Heteropycnosis or precocious staining of part of the nucleus, usually in the form of an open ring, was visible at early prophase of meiosis. However, it was much less pronounced than in many other mosses e.g., Brywm, Rhizogonium, 222 INTRASPECIFIC POLYPLOIDY IN HYPOPTERYGIUM ROTULATUM etc. (Ramsay, unpubl.). Even in dioecious species no evidence was found of a large bivalent which separated precociously and could be related to the possible sex chromosomes described in some other mosses ¢.g., Macromitrium (Ramsay 1966), Pogonatum (Yano, 1955, 1957), ete. Meiosis in spore mother cells of Hypopterygium rotulatum X2,700. 1, 8/64 M-I, n=9, note small bivalent; 2-3, 36/65. 2, M-I, n=9, note small bivalent separating precociously ; 3, A-I small chromosome distinct at each end; 4, 8a/64 M-I, n=18, note two small bivalents ; 5, 33a/64 M-I, n=18, note similarity to fig. 1 but twice the number; 6, 8a/64 M-I, secondary association of bivalents; 7—9, 22/64. n=27; 17, multivalents or secondary association of some bivalents; 8, 27 bivalents at M-L; 9, T-I, note 9 lagging univalents or bivalents; 10-11, 8e/64, n=36; 10, M-I, 36 bivalents visible including four small ones; 11, A-I. H. rotulatum (80/64, 33/64, 33a/64, 336/64) n=18 (Text-figs 4-6. Plate VI, 3-4). The first of these populations was collected on the same log with the previous ones, but the other was found about one mile downstream in Happy Valley (Table 1). Both had 18 bivalents at metaphase of meiosis, consisting of 16 similar in size, and two small precociously disjoming ones. Comparison of HELEN P. RAMSAY 223 bivalents in Plate VI, 1-2 (n=9) and Plate VI, 3 (n=18) suggests that the latter is made up of 2x9 chromosomes. The gametophytes of these must therefore be diploid. Although the populations had the same chromosome number, it was noticed that in 8a/64 secondary association with two or more bivalents associated together at metaphase I was common in dividing cells (Plate VI, 3). During meiosis irregularities observed included univalents left out at telophase I and clumping of groups of chromosomes off the metaphase I plate. Chromosome balance was obviously unstable in this group of plants. Oa Q Haplid Game tophyte BASIC 2 X=9 9 x1g UNREDUCED uneeouceD Jn =27 | 3 Ainphidiploid TY MESEORE NT mi Inz27] (Feplud) ee X Gametophy te £ G © SSKo ra Of i & > y Diploid eet se | ean SP9 Ae Tetraplocd N= 36 Gamet ophyte Fic 12 Possible origin of chromosome races in Hypopterygium rotulatum. However, meiosis in 33/64, 33a/64 and 33b/64 was normal and a very small percentage of cells behaved irregularly. All the plants in this chromosome group on further examination were found to be monoecious, the antheridia being borne in separate perichaetia in the axils of leaves just below the arche- gonial ‘ inflorescences ’. The mitotic chromosomes examined by Shimotomai and Koyama (1932) in the synoecious species H. japonicum consisted of 16 large and two small chromosomes, and compare in relative sizes to those in H. rotulatum with n=18. H. rotulatum (22/64, 8d/64) n=ca. 27 (Text-figs 8-9. Plate VI, 5-11). Meiosis in the spore mother cells of the above groups of plants revealed approximately 27 bivalents at metaphase I. Some had been collected from the same log at Mt. Wilson (Table 1). Only an approximate chromosome count could be obtained, for although some cells had n=27 (Text-fig. 8. Plate VI, 6) clumping, irregularities and multivalent formation or secondary association made it difficult to interpret divisions (Plate VI, 7-10). A maximum of 32 including possible univalents or dissociated bivalents was found and none gave a number as high as 36. Some side views of metaphase I showed a regular arrangement of bivalents on the equator, and anaphase I revealed no lagging chromosomes in a number of the spore mother cells, but others (Text-fig. 9. Plate VI, 11) had as many as nine univalents or bivalents left out at end of anaphase I. Normal tetrads with four equal nuclei were produced in some 224 INTRASPECIFIC POLYPLOIDY IN HYPOPTERYGIUM ROTULATUM spore mother cells, although many contained large and small nuclei, or chromo- somes lying loosely in the cytoplasm even after the spores had been differentiated. H. rotulatum (8e/64, 8b/64, 8c/64, 8f/64, 16/64) n=36 (Text-figs 10-11. Plate VII, 1-8). Table 1 shows that some of these plants also came from the same location at Mt. Wilson. Their chromosome number was found to be n=36. Meiosis was almost regular in 8e/64 although some secondary association was noticed at metaphase I. Separation of chromosomes at anaphase I proceeded normally and lag was infrequent. Tetrad formation appeared normal. In other populations behaviour at meiosis was very irregular. Secondary association of bivalents was common (Plate VII, 4-6), bivalents and univalents lay off the metaphase I plate, lag at anaphase I (Plate VII, 7) was frequent and unequal distribution of chromosomes to nuclei at telophase I and telophase II resulted in irregular tetrads with only three nuclei, or with more than four usually unequal in size. It is unlikely that any viable spores could result from such behaviour. Some breakdown at meiosis might have been due to the excessive numbers of chromosomes or bivalents struggling for position on the equator at metaphase, so that groups of chromosomes, unable to manage this successfully, formed themselves into nuclei containing univalents as well as whole bivalents. Heteropycnosis at early prophase in the polyploids was identical with that in the haploids (Plate VII, 1), no comparable increase in amount or number of heteropycnotic regions being observed. COMPARISON OF TAXONOMIC CHARACTERS A summary of the features of the plants examined is set out in Table 2. Photographs of the leaves and cells of the leaves are found in Plates VIII and IX. The leaves illustrated were taken from side shoots between the middle and the base of the shoot. The gametophyte plants used for these measurements bore several sporophytes, at least two having been examined cytologically. Photomicrographs of cells were taken towards the mid-leaf near the end of the nerve. So that comparison of gametophyte characters could be made in plants with different chromosome numbers, it was assumed that the chromosome number of the gametophyte corresponded to the haploid chromosome number (i.e. n=9 or 18, 27, 36, etc.) obtained at meiosis in the attached capsules. Since all capsules on each plant used had the same chromosome number it seemed reasonable to suppose this to be true, in the absence of mitotic counts. Some of the possible variations in the chromosome numbers of gametophytes giving rise to capsules containing the various chromosome numbers are suggested in the discussion, and the need for actual mitotic counts of gametophyte cells has already been emphasized. Some comments on the methods used for obtaining the measurements are described below. Height of Gametophyte plants. The total height included distance from ground level to the apex of the leafy branches but did not include the spor ophyte Leaf size. The average lengths (not including the point) and breadths (in the mid-leaf) of ten leaves and amphigastria were obtained. Cell size. Twenty cells were examined from areas corresponding to the photographs in Plate IX. Spore mother cells and spore size. Diameters of 20 spores and 20 spore mother cells were measured. Since the spore mother cells were squashed, an area in which the cells were evenly spread was chosen for measurements to give more real comparisons. It was not possible to calculate volumes since the diameter of the squashed cells did not correspond to the original diameter. 225 HELEN P. RAMSAY ‘WppeeiIq—q “YyySUeT=] ‘s]jeo Jeyjour osods="p' W's ‘wBiaysesrydure=-yduy 90:0F g-0F + 91-01 COST 0:6 X GG:P 9°¢ X FP PO 4 HO 8:0 XgG0O-T £ 0:¢-8:T 9€ r9/f8 F0:0F G:0F + §1-01 PPT GG X= 8h BG xX L:& G:0 xX G-0 L:O0 X€6:-0 e 0:6-G:T LG 69/26 S0-0F ¥F:0F + I-11 19-T VG Xe IEG OG 6.7. POX YO —AeD << Teil P 8-2-% 81 r9/ee G0-0F &0:0+F VI-Gl 66-0 6 X L:& BS X 8°& G0 XxX G-0 L:0 X O-1 6 pue P G:°6-G 6 49/8 q XT x 7 ra va aN q XT qx] Beets) =u OZIg ‘OWNS ‘ydury jeoT “UU “WUE Keg eyAydoyoures Tequm N requin eaodg “WRI, ezig -ydury ZIG JRo'T jo 4yusIOFT moo Conae (@) reyono A 9218 190 a ee ee ee ee ee ee ee a a ee a ee wnyzepnjor umiskieydodAy fo sorsrsazonunyo yoorbojoyduow fo uosruodwog @ WIavy, 226 INTRASPECIFIC POLYPLOIDY IN HYPOPTERYGIUM ROTULATUM The results set out in Table 2 give some indication of the features most likely to be of use in further investigations, particularly of gametophytes from which mitotic counts could be obtained. Although the leaves and amphigastria showed little difference in gross size in the various groups of plants studied, the leaf cells in particular showed increase in length and greater variation in plants with chromosome numbers of n=18 and 36. In Plate IX the numbers of cells in a given area appear to be greater in the haploid and triploid and fewer but larger in the diploid and tetraploid. It has been pointed out by Lewis (1961) that decrease in numbers of cells may account for apparently little change in overall size of some polyploids. The length of setae and size of capsules was similar in all chromosome races, and measurements were not therefore included in Table 2. However, a clear difference in the sizes of spore mother cells can be seen in Table 2, and by comparison of the Text-figures and Plates of the various chromosome forms. Spore size data shows a greater variation in the sizes of spores in polyploids. It is interesting to note that the characters examined in plants with a chromo- some number of n=ca. 27 (22/64) were much closer to the ‘ original haploid ’ (i.e. plants with n=9) than to the others. It appears, therefore, that increase in the length of the cells of the leaf, an increase in the size of spore mother cells and a greater range of variation in the size of spores are features which reflect the increase in chromosome numbers in the various plants. Other characters remained much more stable throughout the groups. DISCUSSION Polyploidy in Mosses. Before chromosome numbers in mosses had derieralle been investigated polyploid gametophytes were produced experimentally by apospory (E. and Em. Marchal, 1911) in Brywm caespiticium and B. argentewm. Later Wettstein (1923-1924, 1928) produced a series of polyploid gametophores in Bryum by treatment of the protonema, or of the spore mother cells at meiosis in Funaria. Auto- and amphidiploid races were produced in several mosses e.g. Funaria hygrometrica n=14 to 28 and 56; the polyploid population of Bryum caespiticium n=10 was called B. corrensii with n=20; Amblystegium serpens from n=12 to 24; and Physcomitrium pyriforme from n=18 to 36 and 72. Increasing numbers of naturally occurring inter- and intra specific polyploids are being reported as the chromosome numbers of mosses from many parts of the world become known (Wylie, 1957; Steere and Anderson, 1954; Khanna, 1960; Yano, 1957 a, b, ¢, etc.). Examples include Atrichum undulatum (Hedw.) P. Beauv. (Lowry, 1954) with n=7, 14 and 21, Mnium spp. (Lowry 1948) n=6, 12 Tortula princeps De Not. n=12, 24+1 and 36-+2 (Steere, 1954; Steere ef al., 1954) Philonotis socia Mitt. (Yano, 1957a) with n=6 and 12 and Octoblepharum albidum Hedw. with n=13 and 26 (Khanna, 1960). They represent species from many families. Octoblepharum albidum was analyzed by Khanna (1960) who found no qualitative differences between the plants in external characteristics such as leaf size, size of leaf cells, etc. Some differences in spore size were noted, although overlapping occurred, and from his drawings it appears that the spore mother cells were larger in the polyploid. The majority of the examples of intraspecific polyploidy represent different populations in different localities e.g. Tortula mucronifoia n=12 and 24, Drepanocladus uncinatus n=12 and 24, Distichium capillaceum n=14 and 28 (Anderson and Crum, 1958). The spore mother cells of the latter were found to be larger in the diploid, although the spores were similar in size. One particularly interesting series of interspecific polyploids exists in the genus Bryum, where the chromosome numbers of n=10, 20, 30, 40 and 50 were discovered in Alaskan species by Steere (1954). HELEN P. RAMSAY 227 Lewis (1961) has summarized the work of Wettstein (1940) on the relation- ship of cell size and volume in artificially induced auto- and allo-polyploids. ‘‘ Generally an increase in size of cells, organs and individuals follows recently induced polyploidy but the extent of such increases varies. The effect is not uniform over the entire polyploid range, cell size usually decreasing, or ceasing to increase especially in the more hybrid polyploids ” (Lewis, loc. cit.). In the auto-polyploids of Physcomitrium pyriforme cell size was shown to increase with increase in chromosome number, particularly in the leaf. A negative correlation was found between the initial number of chloroplasts and their number, following doubling of the chromosome numbers (Lewis, 1961). The number of cells in the polyploid organism may be less, so that increase in cell size may not lead to an increase in the size of the organ (Darlington from Lewis, 1961). A significant feature of size relations of polyploids to normal plants is that recently produced polyploids may show distinct variations but natural polyploids do not. Wettstein (1937) observed that in an artificial diploid strain of Bryum caespiticium which he called B. corrensi, the size of cells and other morphological characters were much larger in the newly produced polyploid, but gradually decreased over a period of 11 years until they had reverted to the size of the original haploid cells ete. During the same period meiosis and spore production also became normal, although very irregular in the newly produced diploid. The stabilization of cell size and other characters and the establishment of normal Spore production were found in the original plants after 11 years, in clones produced vegetatively during this time, and in successive generations produced from the first fertile spores. Khanna (1960) suggested that polyploid populations of Octoblepharum albidum were in a state of reversion to haploid size since variations in the cell size, Spore size etc. between the diploid and haploid plants were not constant. Meiotic behaviour in auto-polyploids of maize, studied in detail by Giles and Randolph (1951), showed a reduction in multivalent frequency at meiosis in successive generations, until within 10 years meiotic behaviour had reverted to normal. Apart from influencing the autosome number, polyploidy must also affect sex balance in dioecious species, whether sex is determined by a sex chromosome or a gene complex on autosomes. Doubling of the chromosome number can occur in several ways in mosses such as regeneration from gametophyte cells in which breakdown of mitosis doubles the chromosome number, by apospory from cells of the seta or capsule, or by reconstituted nuclei in spore production at meiosis. The latter would result in the production of monoecious plants. It has been found generally that polyploids are monoecious (Mehra and Khanna, 1961; Anderson, 1964, for summaries) and apospory is probably the most important factor in the evolution of polyploidy in mosses. The chromo somes of such polyploids would have more affinity for their exact replicas than related chromosomes which may bear other alleles, and secondary association of bivalents might occur more frequently than multivalent formation. Polyploidy in Hypopterygium rotulatum. The area from which the plants of this species was collected has been re-visited at frequent intervals in an attempt to repeat the results, and to study the whole population in greater detail. Unfortunately, 1965 and 1966 have been drought years with unusually low rainfalls, particularly in the summer months. In 1965 plants did not regenerate in large numbers, although protonema and old and immature plants were present in the winter months. Very few capsules were produced, and the only chromosome number obtained was n=9 in a few capsules in September (earlier investigations had been carried out in April, 1964). The population is again showing considerable growth and arche- gonia and antheridia are being produced. Dioecious and monoecious forms are 228 INTRASPECIFIC POLYPLOIDY IN HYPOPTERYGIUM ROTULATUM present, although only a few capsules have been found in 1966, but meiosis has not been detected. Drought could again influence fertilization in these plants. The population is obviously maintained by vegetative means. The necessity for water to carry the sperm to the ovum, especially in dioecious species, is a controlling factor for fertilization. Monoecious forms which allow self fertilization would be able to overcome this if a thin film of water were present, and the antheridia and archegonia were placed close together and ripened simultaneously. The genetic disadvantage of selfing is the establishment of strict homozygosity. Isolating mechanisms such as protandry, or self incom- patibility with their genetic advantages of outbreeding, are disadvantages if fertilization depends rigidly on sufficient water at the correct time. Taxonomic descriptions of H. rotulatum state that it is dioecious, whereas the majority of plants examined in this study were found to be monoecious. The populations which showed almost normal meiosis such as 33/64, 8¢/64 and also produced normal looking spores may have resulted from self fertilizations. Cross fertilization in an area where such a variety of chromosome numbers were present would be particularly hazardous. Some of the possible crosses within such a population are set out in Table 3. TABLE 3 Egg Sperm Expected meiotic behaviour n=9 9 x 9 normal pairing 9 x 18 9 pairs, 9 univalents, or trivalents 9 x 27 9 pairs, 18 univalents or multivalents according to affinities 9 x 36 =©69_ pairs, 27 multivalents or univalents n=18 18 x 18 18 pairs, normal behaviour 18 x 9 9 pairs, 9 univalents or multivalents The ovum may have some control over which sperm are acceptable. Other controls may be affected by the genetic affinities of the two nuclei. In some plants sporophytes reached the stage of differentiation into capsules, or almost to formation of the spore mother cells, then shrivelled up. No environmental factor causing this could be found, as such plants were growing among those with well-developed capsules. They may have been the result of irregular fertilizations. Wettstein (1940) mentions that Bryum corrensw did not produce sporophytes, which matured to form capsules and spores, for several years. During the first few years after the development of this experimental polyploid, embryo sporo- phytes developed only for a short time and capsules were not formed. Within 11 years normal sporophytes were regularly produced. From these studies it is clear that polyploid gametophytes must exist among the haploid plants of Hypopterygium rotulatum, although mitotic counts are needed for confirmation. A scheme setting out the possible origins of the various chromosome numbers is set out in Text-figure 12. Evidence for such a scheme comes from the apparent duplication of complements in plants with n=9 and those with n=18 most obvious in the doubling of the number of minute bivalents. The four small bivalents in plants with n=36 (8e/64) points to a further doubling from n=18. All the polyploids are monoecious, while haploids are dioecious, aS would be expected if the new numbers had resulted from failure of reduction in spores at meiosis. Plants with a chromosome number of n=27 could result from fusion of gametes with chromosome numbers of n=9 and 18, followed by reconstitution at meiosis, giving rise to wnreduced spores HELEN P. RAMSAY 229 The results of this investigation appear to be the first in which a population of mosses growing in close proximity show such high grades of intra specific polyploidy. The levels of polyploidy correspond to tetraploids, hexaploids, and octoploids in Angiosperms. Acknowledgements I am indebted to Professor G. K. Berrie who interested me in the cytology of mosses, to Mr. J. H. Willis for the loan of specimens for checking the identification and to Professor S. Smith-White who read the manuscript. References AnveERrson, L. E., 1964.—Biosystematic evaluation of the Musci. Phytomorphology, 14: 27-41. , and Crum, H., 1958——Cytotaxonomia Studies on mosses of the Canadian Rocky Mountains. Nat. Mus. Canada, Bulletin 160: 1-89. Brotuerus, V. F., 1924.—In Engler and Prantl, Die nattrliche Pflanzenfamilien Ed. 2, Vol. 11. Leipzig. GILES, A., ana RaNDOrPE: L. F., 1951.—Reduction of quadrivalent frequency in auto-tetraploid maize during a period of ten years. Amer. Jour. Bot., 38: 12-17. ; Kaanna, K. P., 1960.—The haploid and spontaneous diploid race in Octoblepharum albidum Hedw. Cytologia, 25: 334-341. Lewis, K. R., 1961.—The genetics of Bryophytes. Trans. brit. bryol. Soc., 4: 111-130. Lowry, R. J., 1948.—A cytotaxonomic study of the genus Mniwm. Mem. Torrey bot. Club, 20(2): 1-42. , 1954.—Chromosome numbers and relationships in the genus Aérichum in North America. Amer. Jour. Bot., 41: 410-414. , Marcuat, E., 1920.—Recherches sur les variations numériques des chromosomes dans la série végétale. Mem. Acad. roy. Belg. Cl. Sci., Ser. 2, 4: 3-108. , and Km., 1911.—Aposporie et sexualité chez les mousses III. Bull. roy. Acad. Belg. Cl. Sct. : 750-778. Meura, P. N., and Kanna, K. P., 1961.—Recent cytological investigations in mosses. Fes. Bull. Panjab Univ., 12: 1-29. Ramsay, Helen P., 1964.—The chromosomes of Dawsonia. The Bryologist, 67: 153-162. , 1966.—Sex chromosomes in Macromitrium. The Bryologist. (in press.). Satvspury, G. O. K., 1955.—‘‘ A hand book of the New Zealand mosses”. Roy. Soc. N.Z. Bull., 5: 412. Suimotomar, N., and Koyama, Y., 1932.—Geschlechtschromosomen bei Pogonatum inflexum Lindb. and chromosomenzahlen bei einigen anderen laubmoosen. Jour. Sci. Hiroshima Univ., (B2) 1: 95-101. STEERE, W. C., 1954.—Chromosome number and behaviour in Arctic mosses. Bot. Gaz., 116: 93-133. , 1958.—Evolution and speciation in mosses. Amer. Nat., 42: 5-20. ; , ANDERSON, L. E., and Bryan, Virginia S., 1954.—Chromosome studies on Californian mosses. Mem. Torrey bot. Club, 20: 1—75. Vaarama, A., 1954.—On the characteristics of the spontaneous bivalent race of Funaria hygro- metrica. Arch. Soc. Vanamo, 9: 395-400. ’ WETTSTEIN, F. von, 1923, 1924.—Kreuzungsversuche mit multiploiden moosrassen. vol. Zentralbl., 43: 71-82, 44: 145-168. j , 1924.—Morphologie und physiologie des formwechsels der moose auf genetischer grundlage. I. Z. indukt. Abstamm.—u. Vererb Lehre, 33: 1-236. . , 1928.—Morphologie und physiologie des formwechsels der moose auf genetischer Grundlage. II. Bibl. Genet., Leipzig, 10: 1-216. , 1940.—Experimentelle untersuchungen zum artsbildungs problem. III. Ber. deutsch. bot. Ges., 58: 374-388. Wvurm, Ann. P., 1957.—Chromosome numbers of mosses. Trans. brit. bryol. Soc., 3: 260-278. Yano, K., 1957.—a. Cytological studies on Japanese mosses. I. VFissidentales, Dicranales, Grimmiales, Eubryales. Mem. Fac. Educ. Niigata Univ., 6: 1-31. , 1957.—b. Cytological studies on Japanese mosses. II. Hypnobryales. Mem. Takada Branch Niigata Univ., 1: 85-127. , 957.—c. Cytological studies on Jap se mosses. III Jbid., 1: 129-259. 230 INTRASPECIFIC POLYPLOIDY IN HYPOPTERYGIUM ROTULATUM EXPLANATION OF PLATES VI-IX. Plate vi Hypopterygium rotulatum, meiosis in spore mother cells, x 2,200. 1-2, M-I, n=9, note small bivalent disjomed in 2. 3-4, M-I, n=18. 3, secondary association in 8a/64. 4, 18 bivalents in 33/64 includes two small ones (tiny spots are excess stain). 5-10, n=ca. 27 various stages of meiosis. 5, premetaphase stage before chromosomes are fully contracted, multivalents indicated by arrows. 6, M-I, ca. 27 bivalents showing. 7-8, side view M-I, showing bivalents left off the plate. 9, clumping of chromosomes at M-I. 10, secondary association or multivalents at M-I. 11, late A-I, showing lagging chromosomes, one bivalent in centre not disjoined. Plate vii. Hypopierygium rotulatum, meiosis in spore mother cells of populations with n=36 chromosomes, x 2,200. 1, heteropyenosis in polyploid. 2-3, M-I, n=36, fairly regular arrangement on the metaphase plate. 4-6, M-I, secondary association, clumping in five and six, while a ring is clear in four. 7, T-I, lag of univalents. 8, tetrad of spores, note chromosomes left in cytoplasm. Plate vii. Hypopierygium rotulatum, leaves from plants with different chromosome numbers, x30. 1?’ plants with n=9 (8/64). 2, plants with n=18 (33/64). 3, plants with n=27 (22/64). 4’ plants with n=36 (8//64). Plate ix. Hypopterygium rotulatum, cells of leaves in Plate VIII, x350. 1, cells from plants with n=9 (8/64). 2, cells from plants with n=18 (33/64). 3, cells from plants with n=27 (22/64). 4, cells from plants with n=36 (8//64). HIGHLANDS PEAT OF THE MALAYAN PENINSULA B. R. Hewitt Chemistry Department, University of Malaya, Kuala Lumpur Malaysia [Read 30th November, 1966] Synopsis The factors responsible for the formation of highland peat on the Malayan Peninsula are mentioned together with analytical results for a peat from Gunong Erau, Malaya. The highland peat of the tropics is quite a distinct formation from the peat found on the lowlands. In contrast to the extensive lowland peat (two million acres in the Malayan Peninsula, 40 million acres in Indonesia) highland peat is found in isolated areas on the tops of mountains where the underlying rock is quite acidic. Mohr in 1922 stated after examination of the Hjang-Plateau in Besoeki, Indonesia, ‘‘ Man findet dort merkwurdigerweise Stellen mit einer Art von bleichem Torf bedeckt welche sich halten kann unter Einwirkung der Kombi- nationen von niedriger Temperatur und des Unterwasserstehens.”’ In 1930 Vageler maintained that only those mountain peats formed from Sphagnum were genuine Highmoors. This would appear to be too rigid a classification. Tropical highmoors contain Sphagnum moss but in Malaya, at least, there is also present forest vegetation which, however, does not reach the full development of the lowland forest and does contribute material for the formation of peat. In Indonesia, highmoor peats .are formed in Sumatra, Toba-Ebene, 1,450 m. (4,750 ft.) ; in Java, at Gedeh, Telaga-Saat, 1,400 m. (4,570 ft.) , Dieng Plateau, 2,000 m. (6,500 ft.) ; in Celebes, at Lindessee, 1,000 m. (3,260 ft.). Many of the craters of Indonesian mountains contain swamps in which peat forms. In addition to Sphagnum and Juncus prismatocarpus R.Br., swamp grasses and Cyperaceae grow on the banks (von Faber, 1927). This type of peat present in the tropics is found in many parts of the world and is typical of Europe and other areas with temperate climates. They occur in Australia in humid localities, usually in parent materials of low base status and the vegetation is often dominated by Cyperaceae (Stephens, 1949-50). On the Malayan Peninsula highmoor peat is formed at altitudes greater than 1,500 m. (4,900 ft.), usually on relatively bare quartzite or acid granite ridges (Scrivenor, 1931) having a typical vegetation of montane ericaceous forest with sphagnum and lichens on the ground (Wyatt-Smith, 1954). The montane ericaceous forest occurs above 1,500 m. (4,900 ft.) on exposed ridges and consists of dwarfed vegetation carrying masses of epiphytes. The main species present are: FAGACEAE, Pasania lampadaria, Pasania rassa ; ERICACEAE, Pieris ovalifolia, Vaccinium spp. Rhododendron spp.; HELAEOCAR- PACEAE, Hlaeocarpus masterii ; MYRTACEAE, Hugenia spp., Rhodamnia cinerea, Tristania merguensis; THEACEAE, Ternstroemia japonica, Anneslea crassipes ; SYMPLOCACEAE, Symplocos spp.; KUTACEAE, Astrophyllum montanum, Tetrac- tomia tetrandra; GUTTIFERAE, Garcinia spp.; AQUIFOLIACEAE, Ilex spp. ; MYRSINACEAE, Myrsine posteriana; ULAURACEAE, Phoebe declinata; PENTA- PHYLACACEAE, Pentaphylax arborea; BUXACEAE, Buxus spp. Peat forms a layer on the ground to a maximum depth of about 5 cms. but will accumulate to greater depths in rock crevasses. It is not associated PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wates, Vol. 91, Part 3 232 HIGHLANDS PEAT OF THE MALAYAN PENINSULA with swamps or ponds, and its accumulation is due to the continually humid conditions and low temperatures preventing oxidation of the plant material and to the presence of very acid rock underlying it. Samples of peat were taken from Gunong Erau in the Cameron Highlands at the height of approximately 1,960 m. (6,400 ft.). Here the daily average temperature ranges from 14-1-23°C. (57-6-73-4°F.), the relative humidity at 1 p.m. is greater than 98°, throughout the year, while the total rainfall is 2,687 mm. (105-5 inches) and the lowest average rainfall in any month is 129 mm. (five inches). There was no visible difference in the samples which were of a reddish brown colour. The results of analysis are given in Table 1. TABLE | Analysis of peat from Gunong Hrau Sample pH. Ash content % Loss on ignition % depth (on dry basis) (on dry basis) 0-20 em. (0-8 inches) 2-0 2-6 97-4 20-40 cm. (8-16 inches) 2-0 2-3 Q7°'7 40-60 em. (16—24 inches) .. 2-3 ToB 92-7 The pH was determined on dried peat in IM.KCl with a soil KCl ratio of 1 to 5. Humus fractions were determined according to the procedure of Konanova, 1961, and are given in Table 2. TABLE 2 Humus fraction (Percentage of dry weight of sample) Sample Humin Hymato Crenic + Humic + depth + -melanic Apocrenic Ulmic Ulmin acid acids acids 0-20 em. (0-8 inches) .. 25-0 14-6 55-4 5:0 20-40 em. (8-16 inches) 30-0 10-4 53-0 7-3 40-60 em. (16—24 inches) 25-0 18-5 47-5 9-0 The low ash contents and pH’s of all the samples are striking. There is a slight decrease in acidity and increase in ash content with increase in depth. Analysis of similar material at the University of Liverpool (Burges, 1966) showed that it contained degradation products characteristic of humic acid found under broad-leaved forests in Europe (Burges, et al., 1964). The similarity to peat material from temperate regions is not unusual, since the climate of the Cameron Highlands of Malaya is temperate and the vegetation is similar to heath vegetation. References Burass, N. A., Hurst, H. M., and WAKKENDEN, B., 1964.—The phenolic constituents of humic acid and their relation to the lignin of the plant cover. Geochem. et Cosmochim. Acta. 28: 1547-1554. Burcss, N. A., 1966.—Private communication. Fapur, F. C., von, 1927.—‘‘ Die Kraterpflanzen Javas Welterreden ”’. Konanova, M. M., 1961.—Soil Organic Matter. (Pergamon Press, Oxford.). Monr, EH. C. J., 1922.—** De grond van Java en Sumatra”. (Amsterdam.). Scrivenor, J. B., 1931.—‘‘ The Geology of Malaya’. (Macmillan, London.). STEPHENS, C. G., 1949-50.—Comparative morphology and genetic relationships of certain Australian, North American and European soils. J. Sol. Sczv., 1(2): 124-129. VaAGELER, P., 1930.—‘‘ Grundriss der tropischen und subtropischen Bodenkunde’’. (Berlin.). Wvatt-Smitu, J., 1954.—Forest types in the Federation of Malaya. Malay Forester, xvii: 83-84. ABSTRACT OF PROCEEDINGS ORDINARY MONTHLY MEETING 30th MARCH, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (24th November, 1965) were taken as read and signed. The following were elected Ordinary Members of the Society: Messrs. A. A. Martin, B.Sce.(Hons.) Rand., University of Melbourne, Parkville, Victoria ; and R. HE. Wass, B.Se.(Hons.) (Q’ld.), Department of Geology and Geophysics, University of Sydney. The Chairman announced that library accessions amounting to 50 volumes, 495 parts or numbers, 15 bulletins, 13 reports and 10 pamphlets, total 583, had been received since the last meeting. The Chairman drew the attention of members to a Seminar on The Bush Fire Problem in Public Lands to be conducted at Leura on 16th and 17th April under the auspices of the National Trust of Australia (N.S.W.). PAPERS READ (By title only, an opportunity for discussion to be given at the April Ordinary Monthly Meeting) 1. The composition, occurrence and origin of lerp, the sugary secretion of Spondyliaspis eucalyptt (Dobson). By R. Basden. 2. Seeds and fruit of the Goodeniaceae. By R. C. Carolin. 3. The breeding biology and larval development of Hyla jervisiensis (Anura : Hylidae). By A. A. Martin and M. J. Littlejohn. ORDINARY MONTHLY MEETING 27th APRIL, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (30th Mareh, 1966) were read and confirmed. The Chairman announced that the Council had elected the following office-bearers for the 1966-67 session: Vice-Presidents: Dr. D. T. Anderson, Miss Elizabeth C. Pope, Mr. G. P. Whitley and Professor J. M. Vincent ; Honorary Treasurer: Dr. A. B. Walkom; Honorary Secretary, Mr. R. H. Anderson. The Chairman referred to the retirement of Dr. W. R. Browne from the position of Honorary Secretary and to his great services to the Society in this and other capacities. 234 ABSTRACT OF PROCEEDINGS The Chairman announced that library accessions amounting to 15 volumes, 158 parts or numbers, 1 bulletin, 1 report and 1 pamphlet, total 176, had been received since the last meeting. The Chairman reminded members that there would be no Ordinary Monthly Meeting in May. PAPERS READ 1. Fungi attacking seeds in dry seed-beds. By D. M. Griffin. 2. On the species Fenestella horologia Bretnall and Minilya duplaris Crockford. By R. Wass. 3. Two new species of Permian Brachiopods from Queensland. By BR. Wass, HXHIBIT Mr. A. G. Khan (a visitor introduced by Dr. I. V. Newman) exhibited embryos of Podocarpus species grown in agar culture. Fully grown embryos, excised under aseptic conditions from the surface-sterilised seeds, were cultured aseptically on two culture media differing chiefly in that one had coconut milk and a greater number of growth-factors. The culture tubes were kept at room temperature. Dry P. elatus R. Br. seeds were supplied by the Forestry Commission (from low-temperature storage). The embryos from these seeds were cultured on 26th March last and are still dormant on April 27, not Showing any growth on the surface of either medium. Full size green P. faleatus R. Br. seeds were collected from the Royal Botanic Gardens, Sydney, on 14th April, 1966, and their embryos were cultured on the same day. By 27th April the embryos are showing significant growth on the surface of both media, the cotyledons having turned green and the roots appearing. The embryos from the same collection of P. falcatus R. Br. seeds when submerged in either of the two culture media remained dormant. LECTURETTE Miss Elizabeth C. Pope, Australian Museum, Sydney, gave a lecturette on the fauna of the intertidal reefs at Darwin, Northern Territory, illustrated by colour transparencies. ORDINARY MONTHLY MEETING 29th JUNE, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (27th April, 1966) were read and confirmed. ' The following were elected Ordinary Members of the Society: Dr. John Child, M.A., B.Comm. (N.Z.), D.Phil. (Oxon.), Gladesville, N.S.W.; Messrs. G. A. Holloway, Australian Museum, Sydney; and A. G. Khan, Surry Hills, N.S.W. The Chairman announced that Dr. C. G. Hansford, who had been a member of the Society since 1952, had died at Uvongo Beach, Natal, South Africa, on 18th February, 1966. ABSTRACT OF PROCEEDINGS 235 The Chairman announced that library accessions amounting to 31 volumes, 301 parts or numbers, 10 bulletins, 7 reports and 6 pamphlets, total 355, had been received since the last meeting. The Chairman announced that the next Ordinary Monthly Meeting will be held in the Main Hall of Science House on Friday, 15th July, at 7.45 p.m., immediately preceding the Sir William Macleay Memorial Lecture to be delivered at 8 p.m., by Professor D. G. Catcheside, D.Sc., F.R.S., the subject being ‘‘ The Centenary of Mendel ”’. The Chairman also announced that there will be no Ordinary Monthly Meeting in August. The Chairman referred to the retirement of Dr. A. B. Walkom as Honorary Secretary and to his valuable services to the Society over many years. A vote of thanks to Dr. Walkom was carried by acclamation. The Chairman announced the appointment of Dr. M. F. Day as a Trustee of the Kosciusko State Park and referred to the satisfaction of the Society at such action. PAPERS READ 1. The Rhaphidophoridae (Orthoptera) of Australia. Part 4. A new genus from South Australia. By Aola M. Richards. 2. Notes on three recently proposed Australian Tertiary echinoid genera. By G. M. Philip. 3. Chromosome numbers in Lamprothamnium. By A. T. Hotchkiss. 4, Mosquitoes of Tasmania and Bass Strait Islands. By N. V. Dobrotworsky. NOTES AND EXHIBITS Mr. G. P. Whitley contributed a note on announcements of the formation of the Linnean Society of New South Wales in The Sydney Morning Herald newspapers of October and November, 1874. On 30th October, 1874, page 4, appeared : ‘SCIENCE OF NATURAL HISTORY.—A preliminary meeting of gentlemen interested in the study of natural science was held yesterday in the boardroom of the Free Library, and, after some discussion, the following resolutions were carried unanimously :—‘ That a society for the cultivation of the science of Natural History in all its branches be now formed, under the name of ‘‘ The Linnean Society of New South Wales”, and that all those whose names are signed to the papers now before the meeting shall be, and are, the original members of such society.—That the annual subscription shall be £1 1s., payable on the 1st January of each year; and that all members joining after the close of the present year shall pay an entrance fee of £1 1s., in addition to their annual subscription.—That the officers of the society shall consist of a president, vice- president, secretary, treasurer, and a council of six.—That the following gentlemen be requested to accept the respective offices :—President, Mr. Macleay ; vice-president, Sir W. Macarthur; secretary, Captain Stackhouse; treasurer, Mr. Burton Bradley.—That a committee, consisting of the president, Mr. Stephens, and Dr. Alleyne, be appointed to draw up rules to lay before the adjourned meeting this day week, to make inquiries respecting a room for the use of the society, and generally to have full powers to conduct the affairs of the society until the election of the Council.—That the above-mentioned committee be instructed to draw up and distribute, according to their discretion, a circular, giving full information as to the nature and aims of the society, and inviting the co-operation of all interested in the study of natural history —That a concise account of the proceedings of this meeting be forwarded to the Sydney daily newspapers, and that an advertisement be inserted in each, stating the time and place of the next meeting.’ The meeting was then adjourned till next week.” ¥F 236 ABSTRACT OF PROCEEDINGS The above paragraph was repeated in the Herald of 31st October, page 6. On page 5 of the 5th November issue, the Herald announced that the Linnean Society of New South Wales was to meet that afternoon to consider draft rules, and there is a brief account of the meeting in The Sydney M Cao Herald, 6th November, 1874, page 5. Newspapers have been consulted at the Public Library of New South Wales. LECTURETTE An illustrated lecturette entitled ‘‘ Plants in Iceland ” was given by Dr. D. J. Anderson, School of Biological Sciences, Department of Botany, University of Sydney. ORDINARY MONTHLY MEETING 15th JuLy, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (29th June, 1966) were taken ag read and confirmed. The Chairman announced that library accessions amounting to 13 volumes, 185 parts or numbers and 3 bulletins, total 201, had been received since the last meeting. The Chairman announced that there will be no Ordinary Monthly Meeting in August. PAPERS READ (By title only, an opportunity for discussion to be given at the September Ordinary Monthly Meeting) 1. The application of the names Sericesthis pruinosa (Dalman) and Sericesthis nigrolineata Boisduval (Coleoptera: Scarabaeidae: Melolonthinae). By E. B. Britton. (Communicated by Dr. D. F. Waterhouse.) 2. New taxa of Acacia from eastern Australia. No.2. By Mary D. Tindale. ORDINARY MONTHLY MEETING 28th SEPTEMBER, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (15th July, 1966) were read and confirmed. The following were elected Ordinary Members of the Society: Mrs. Denise D. Clyne, Turramurra, N.S.W.; Mr. B. A. Conroy, Leichhardt, N.S.W. ; Miss Judith H. Ford, Mosman, N.S.W. ; Miss Lois J. Lovedee, B.Se. , East Gosford, N.S.W.; Mr. Ahmed Mahmood, M. 'Se., University of Karachi, Karachi- 32, West Pakistan ; and Mr. G. L. Miklos, Hunter’s Hill, N.S.W. The Chairman announced that Mr. George N. Baur had been elected a member of Council in place of Professor 8S. Smith-White. ABSTRACT OF PROCEEDINGS 237 The Chairman announced that the Council is prepared to receive applications for Linnean Macleay Fellowships tenable for one year from 1st January, 1967, from qualified candidates. Each applicant must be a member of this Society and be a graduate in Science or Agricultural Science of the University of Sydney. The range of actual (tax-free) salary is, according to qualifications, up to a maximum of $A3,200 per annum. Applications should be lodged with the Honorary Secretary, who will give further details and information, not later than Wednesday, 2nd November, 1966. The Chairman announced that library accessions amounting to 39 volumes, 318 parts or numbers, 19 bulletins, 16 reports and 17 pamphlets, total 409, had been received since the last meeting. PAPERS READ 1. Studies on the inheritance of rust resistance in oats. IV. Expression of allelomorphism and gene interaction for crown rust resistance in crosses involving the oat variety Bond. By EH. P. Baker and Y. M. Upadhyaya. 2. A new corallanid isopod parasitic on Australian freshwater prawns. By EK. F. Riek. 3. New Australian cave Carabidae (Coleoptera). By B. P. Moore. LECTURETTE A lecturette entitled ‘‘ Dolphins and small whales in the Solomon Islands and New Guinea waters’, illustrated by still and movie films, was delivered by Dr. W. H. Dawbin, School of Biological Sciences, University of Sydney. ORDINARY MONTHLY MEETING 26th OCTOBER, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (28th September, 1966) were read and confirmed. Mr. B. F. Clough, B.Sc.Agr., Sydney University; Mr. 8S. W. L. Jacobs, Pymble, N.S.W.; Dr. Patricia Mather (née Kott), University of Queensland, St. Lucia, Queensland; Mr. J. W. Patrick, B.Sc.Ag., Sydney University ; and Miss Hileen F. Pyne, B.Sc., Strathfield West, N.S.W., were elected Ordinary Members of the Society. The Chairman announced that the Council is prepared to receive applications for Linnean Macleay Fellowships tenable for one year from 1st January, 1967, from qualified candidates. Each applicant must be a member of this Society and be a graduate in Science or Agricultural Science of the University of Sydney. The range of actual (tax-free) salary is, according to qualifications, up to a maximum of $A3,200 per annum. Applications should be lodged with the Honorary Secretary, who will give further details and information, not later than Wednesday, 2nd November, 1966. The Chairman announced that library accessions amounting to 18 volumes, 135 parts or numbers, 3 bulletins, 4 reports and 2 pamphlets, total 162, had been received since the last meeting. 238 ABSTRACT OF PROCEEDINGS PAPERS READ 1. Atopozoa deerata (Sluiter): a discussion of the relationship of the genus and species. By Patricia Kott (Dr. Patricia Mather). 2. Inheritance of resistance to bunt in Turkey wheat selections. By E. P. Baker. . LECTURETTE An illustrated lecturette entitled ‘‘ Phylogeny and Evolutionary Patterns in the Flowering Plant Family Dipsacaceae”’ was delivered by Professor F. Ehrendorfer, Director, Institute of Systematic Botany, and Botanic Gardens, University of Graz, Austria. ORDINARY MONTHLY MEETING 30th NOVEMBER, 1966 Dr. R. C. Carolin, President, in the chair. The minutes of the last Monthly Meeting (26th October, 1966) were read and confirmed. The following were elected Ordinary Members of the Society: Mr. R. A. Boyd, B.Sc., University of New England, Armidale, N.S.W., and Mr. EH. H. Hoult, B.Se.(Hons.), University of New England, Armidale, N.S.W. The Chairman announced that the Council had appointed Miss Alison K. Dandie, B.Sc.(Hons.), to a Linnean Macleay Fellowship in Botany for one year from 1st January, 1967. The Chairman referred to the death on 6th October, 1966, of Mr. Melbourne Ward, who had been a member of the Society since 1930. The Chairman announced that library accessions amounting to 19 volumes, 248 parts or numbers, 14 bulletins, 23 reports and 9 pamphlets, total 313, had been received since the last meeting. PAPERS READ 1. Rhinonyssine nasal mite infestations in birds at Mitchell River Mission during the wet and dry seasons. By R. Domrow. 2. Intraspecific polyploidy in Hypopterygium rotulatum (Hedw.) Brid. By Helen P. Ramsay. 3. Highlands peat of the Malayan Peninsula. By B. R. Hewitt. NOTES AND EXHIBITS Mr. Ellis Troughton (Hon. Science Associate at the Australian Museum) exhibited the skull of the adult ¢ holotype of the New Guinea Highland Dog, Canis hallstromi (Proc. roy. zool. Soc. N.S.W., May, 1957), from the Huri-Duna country, Southern Highlands district of Papua. Received at Taronga Park Zoo in March, 1957, the original pair produced a series of litters, invariably black at birth, resulting in some variability in adult coloration. There are several references to a small native dog in New Guinea with a yodelling call, from as far back as 1842 in the Colonial Magazine. The first skin and skull to reach the Queensland Museum were collected by the Lieut. Governor, Sir Wm. MacGregor, in 1897, at 7,000 ft. on Mt. Scratchley in Papua. ABSTRACT OF PROCEEDINGS 239 The specimen was the subject of a rather misleading colour description, and unduly dingo-like photo of the stuffed skin, provided by De Vis (Ann. Qld. Mus., 10, 1911). Subsequently Professor Wood Jones (American Journ. Mamm., X, 1929) published a description and figure of the skull, concluding that it provided definite evidence ‘‘ that the Papuan Dog is a very definite race, possessing a relatively large upper carnassial tooth .. . and differing widely in its characters from the dogs of certain other Pacific Islands ’’. He also stressed the need for study of additional material before the breed became too hybridized. Wood Jones did not confirm his diagnosis with a specific or racial name so that his paper has been generally overlooked. ‘Therefore, in view of the persistence of such distinctive characters in the original pair and their progeny - at Taronga Park, confirmed by personal examination of the skull and skins in the Queensland Museum, the unusual course of specifically naming the living pair, within two months of arrival at Taronga Park, obviated the inevitable inference that the progeny were subject to captive hybridization. The term ‘ feral ’’? was used in Wood Jones’ racial definition, but the term may be applied too casually in discrediting the New Guinea Highland Dog as a mere variation of the mainland Dingo, currently estimated as existing for about 22,000 years in its present form. More probably, the New Guinea Dog was a prehistoric inhabitant of the rain-forests of the Atherton Tableland and, by subsequent deployment over the more arid regions, became the actual progenitor of the Australian Dingo. Mr. A. G. Khan exhibited photographs and photomicrographs concerning the nodulation of roots in three species of Podocarpus. Photographs showed the rooted cuttings bearing nodules in sterile culture of P. elatus R.Br. ex End., P. spinulosus (Sm.) R.Br. ex Mirb., and P. lawrencet Hook f., the last mentioned one being the most prolific in nodule and adventitious root formations. Phyto- micrographs showed the development of nodule and lateral root on naturally growing infected plants. The cortical tissue of the nodule produced in sterile culture is devoid of any endophyte which eliminates infection as a cause of the nodulation. This is strengthened by the anatomical features of young nodule and young lateral root, both of which arise endogenously but differ in their cellular configuration. The nodule lacks any appearance of an apical meristem pattern of cells and has an endodermis over-arching its vascular tissue, whereas the lateral root shows an apical meristem pattern of cells and has an open-ended endodermis. Prof. T. G. Vallance exhibited his copy of George Shaw’s “‘ Zoology of New Holland ” (London, 1794) and a MS letter from Shaw to the artist James Sowerby giving instructions for an illustration in the book. The letter (undated) contains the following passage : ‘“‘ Dr. Shaws Compts. to Mr. Sowerby & informs him that the quadruped intended for the ensuing No. of the New-Holland Zoology is now at Mr. Wilson’s, & if Mr. S. will send a messenger for it he may have it at his own house for some days, which will be necessary, in order to study its several attitudes, & to give as elegant a figure of it as possible. It is an Opossum with the aspect of a Squirrel, & is a very beautiful animal... Mr. S. will take notice that the tail is strongly prehensile, & may therefore be represented in such a manner as to shew that particular, unless it shd. be thought to interfere with the elegance of the plate; But the best way will be to make several sketches, in different attitudes. It is to be fed with bread & milk. It is nearly torpid by day, but very active by night. Care must be taken to express well & clearly the lateral membrance of the sides & feet, as in the flying Squirrel...” The animal referred to in the letter appears in Tab. XI of Shaw’s book, where it is identified as Didelphis sciurea, the Squirrel Opossum. 240 ABSTRACT OF PROCEEDINGS Dr. R. C. Carolin showed a film, prepared by himself and Mr. R. J. Oldfield, demonstrating pollinating mechanisms in Australian native plants. It showed pollination by birds and insects in the families Proteaceae, Myrtaceae, Stylidiaceae, Papilionaceae and Dilleniaceae. Numerous insects were shown working flowers in their own distinctive ways, some clearly pollinating flowers, whereas the insects of others could have had no effect in this direction. The explosive mechanism of Conospermum was illustrated. Mr. 8S. W. L. Jacobs exhibited photographs illustrating the distribution of starch in transverse sections of fresh grass leaves. Podideae show general starch distribution, whereas in Panicoideae and Eragrostoideae starch formation is restricted to the parenchymatous sheath around the vascular bundle. In Triodia, where the photosynthetic tissue is more restricted, this starch-forming sheath surrounds the chlorenchyma rather than the vascular bundle. It is suggested that the parenchymatous sheath is, functionally, more connected with the chlorenchyma than with the conducting tissue. 1940 1940 1965 1959 1964 1922 1963 1927 1952 1912 1962 1961 1949 1962 1959 1950 1960 1955 1965 1960 1954 1935 1946 1940 1964 1961 LIST OF MEMBERS. 241 LIST OF MEMBERS. (15th December, 1966) ORDINARY MEMBERS. (An asterisk (*) denotes Life Member.) Abbie, Professor Andrew Arthur, M.D., B.S., B.Sc., Ph.D., c.o. University of Adelaide, Adelaide, South Australia. *Allman, Stuart Leo, B.Sc.Agr., M.Sc., 99 Cumberland Avenue, Collaroy, N.S.W. Anderson, Derek John, Ph.D., School of Biological Sciences, Botany Building, Sydney University. Anderson, Donald Thomas, B.Sec., Ph.D., School of Biological Sciences, Department of Zoology, Sydney University. Anderson, Mrs. Jennifer Mercianna Elizabeth, B.Sc.Agr., Macleay Museum, Sydney University. Anderson, Robert Henry, B.Sc.Agr., 19 Kareela Road, Chatswood, N.S.W. Ardley, John Henry, B.Se. (N.Z.), School of Public. Health and Tropical Medicine, Sydney University. *Armstrong, Jack Walter Trench, ‘‘Cullingera’, Nyngan, N.S.W. Ashton, David Hungerford, B.Sc., Ph.D., 92 Warrigal Road, Surrey Hills, #.10, Victoria. Aurousseau, Marcel, B.Sc., 229 Woodland Street, Balgowlah, N.S.W. Bailey, Peter Thomas, B.Sc., C.S.I.R.O., Division of Wildlife Research, P.O. Box 109, City, Canberra, A.C.T. Bain, Miss Joan Maud, M.Sc., 18 Onyx Road, Artarmon, N.S.W. Baker, Eldred Percy, B.Sc.Agr., Ph.D., Department of Agricultural Botany, Sydney University. Ballantyne, Miss Barbara Jean, B.Sc.Agr., N.S.W. Department of Agriculture Private Mail Bag No. 10, Rydalmere, N.S.W. Bamber, Richard Kenneth, F.S.T.C., 113 Lucinda Avenue South, Wahroonga, N.S.W. *Barber, Professor Horace Newton, M.S., Ph.D., F.A.A., School of Biological Sciences, Department of Botany, University of N.S.W., P.O. Box 1, Kensington, N.S.W. Barber, Ian Alexander, B.Sc.Agr., School of Biological Sciences, Department of Zoology, Sydney University. Barlow, Bryan Alwyn, B.Sc., Ph.D., School of Biological Sciences, The Flinders University of South Australia, Bedford Park, South Australia. Basden, Ralph, M.Ed., B.Sc. (Lond.), F.R.A.C.I., A.S.T.C., 183 Parkway Avenue, Hamilton, N.S.W. Batley, Alan Francis, A.C.A., 123 Burns Road, Wahroonga, N.S.W. Baur, George Norton, B.Se., B.Se.For., Dip.For., 3 Mary Street, Beecroft, N.S.W. *Beadle, Professor Noel Charles William, D.Sc., University of New England, Armidale, 5N, N.S. W. : Bearup, Arthur Joseph, B.Sc., 66 Pacific Avenue, Penshurst, N.S.W. Beattie, Joan Marion, D.Se. (née Crockford), 2 Grace Avenue, Beecroft, N.S.W. Bedford, Geoffrey Owen, B.Sc., c/- C.S.1.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Bedford, Miss Lynette, B.Sc., School of Biological Sciences, Department of Zoology, Sydney University. Bennett, Miss Isobel Ida, Hon. M.Sc., School of Biological Sciences, Department of Zoology, Sydney University. Bertus, Anthony Lawrence, B.Sc., Biology Branch, N.S.W. Department of Agriculture, Private Mail Bag, No. 10, Rydalmere, N.S.W. Besly, Miss Mary Ann Catherine, B.A., School of Biological Sciences, Department of Zoology, Sydney University. © Bishop, James Arthur, Department of Genetics, The University of Liverpool, Liverpool 3, England. Blackmore, John Allan Philip, LL.B. (Syd. Univ.), 25 Holden Street, Ashfield, N.S.W. Blake, Clifford Douglas, B.Sc.Agr., Ph.D., Faculty of Agriculture, Sydney University. Blake, Stanley Thatcher, D.Sc. (Q’ld.), Botanic Gardens, Brisbane, Queensland. Bourke, Terrence Victor, B.Sc.Agr., c/- Department of Agriculture, Stock and Fisheries, Popondetta, Papua. Brett, Robert Gordon Lindsay, B.Se., 7 Petty Street, West Hobart, Tasmania. Brewer, Ilma Mary, D.Se., 7 Thornton Street, Darling Point, Sydney. Briggs, Miss Barbara Gillian, Ph.D., 13 Findlay Avenue, Roseville, N.S.W. Browne, Ida Alison, D.Sc. (née Brown), 368 Edgecliff Road, Edgecliff, N.S.W. Browne, William Rowan, D.Sc., F.A.A., 363 Edgecliff Road, Edgecliff, N.S.W. Burden, John Henry, 1 Havilah Street, Chatswood, N.S.W. 242 1931 1959 1962 1960 1959 1927 1934 1949 1956 1957 1936 1956 1966 1959 1947 1932 1946 1961 1947 1901 1966 1966 1956 1946 1956 1966 1942 1947 1964 1950 1957 1965 1965 1960 1957 1945 1953 1954 1960 LIST OF MEMBERS. *Burges, Professor Norman Alan, M.Sc., Ph.D., Professor of Botany, University of Liver- pool, Liverpool, England. Burgess, The Rev. Colin E. B. H., Parks and Gardens Section, Department of the Interior, Canberra, A.C.T. Burgess, Ian Peter, B.Sc.For., Dip.For., The Forestry Office, Coff’s Harbour, N.S.W. Cady, Leo Isaac, P.O. Box 88, Kiama, N.S.W. Campbell, Keith George, D.F.C., B.Se.For., Dip.For., M.Sc., 17 Third Avenue, Epping, N.S.W. Campbell, Thomas Graham, Division of Entomology, C.S.I.R.O., P.O. Box 109, City, Canberra, A.C.T. *Carey, Professor Samuel Warren, D.Sc., Geology Department, University of Tasmania, Hobart, Tasmania. Carne, Phillip Broughton, B.Agr.Sci. (Melb.), Ph.D. (London), D.I.C., C.S.I.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Carolin, Roger Charles, B.Sce., A.R.C.S., Ph.D., School of Biological Sciences, Department of Botany, Sydney University. Casimir, Max, B.Sc.Agr., Entomological Branch, N.S.W. Department of Agriculture, Private Mail Bag, No. 10, Rydalmere, N.S.W. *Chadwick, Clarence Earl, B.Sc., Entomological Branch, N.S.W. Department of Agriculture, Private Mail Bag No. 10, Rydalmere, N.S.W. Chambers, Thomas Carrick, M.Sc. (N.Z.), Ph.D., Botany School, University of Melbourne, Parkville, N.2, Victoria. Child, John, M.A., B.Ccomm. (N.Z.), D.Phil. (Oxon.), 48 Milling Street, Gladesville, N.S.W. Chippendale, George McCartney, B.Sc., 4 Raoul Place, Lyons, A.C.T. Christian, Stanley Hinton, Malaria Research Unit and School, Kundiawa, Eastern High- lands, Territory of Papua and New Guinea. *Churchward, John Gordon, B.Sc.Agr., Ph.D., “Shady Trees’, Clarktown Avenue, Mount Eliza, Victoria. Clark, Laurance Ross, M.Sc., c.o. C.S.I.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Clarke, Miss Lesley Dorothy, 4 Gordon Crescent, Hastwood, N.S.W. Clarke, Mrs. Muriel Catherine, M.Sc (née Morris), 122 Swan Street, Morpeth, N.S.W. Cleland, Professor Sir John Burton, M.D., Ch.M., C.B.E., 1 Dashwood Road, Beaumont, Adelaide, South Australia. : Clough, Barry Francis, B.Sc.Agr., School of Biological Sciences, Department of Botany, Sydney University. Clyne, Mrs. Denise Dorothy, 7 Catalpa Crescent, Turramurra, N.S.W. Cogger, Harold George, M.Sc., Australian Museum, College Street, Sydney. Colless, Donald Henry, Ph.D. (Univ. of Malaya), c.o. Division of Entomology, C.S.1.R.O., P.O. Box 109, City, Canberra, A.C.T. Common, Ian Francis Bell, M.A., M.Sce.Agr., C.S.I.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Conroy, Brian Alfred, 22 Carlisle Street, Leichhardt, N.S.W. Copland, Stephen John, M.Sce., 15 Chilton Parade, Warrawee, N.S.W. Costin, Alex Baillie, B.Sc.Agr., C.S.I.R.O., Division of Plant Industry, P.O. Box 109, City, Canberra, A.C.T. Craddock, Miss Elysse Margaret, 36 Lyons Road, Drummoyne, N.S.W. Crawford, Lindsay Dinham, B.Sc., c/- Victorian Plant Research Institute, Department of Agriculture, Burnley Gardens, Melbourne, Victoria. Crook, Keith Alan Waterhouse, M.Sc., Ph.D. (New England), Department of Geology Australian National University, G.P.O. Box 197, Canberra, A.C.T. Croucher, Miss Phillipa Audrey, B.Sc., Department of Zoology, Australian National University, P.O. Box 4, Canberra, A.C.T. Dandie, Miss Alison Kay, B.Sc. (Hons.), 160 Alt Street, Haberfield, N.S.W. Dart, Peter John, B.Sc.Agr., Ph.D., Soil Microbiology Department, Rothamsted Experi- mental Station, Harpenden, Herts., England. Davies, Stephen John James Frank, B.A. (Cantab.), C.S.I.R.O., Private Bag, Nedlands, Western Australia. Davis, Professor Gwenda Louise, Ph.D., B.Sec., Faculty of Science, University of New England, Armidale, 5N, N.S.W. Dobrotworsky, Nikolai V., M.Sc., Ph.D., Department of Zoology, University of Melbourne, Parkville, N.2, Victoria. Domrow, Robert, B.A., B.Sc., Queensland Institute of Medical Research, Herston Road, Herston, N.9, Brisbane, Queensland. Dorman, Herbert Clifford, J.P., A.S.T.C. (Dip.Chem.), Dip.Soec.Stud. (Sydney), Rodgers Street, Teralba 2N. N.S.W. 1954 1946 1952 1953 1947 1930 1957 1964 1955 1957 1964 1966 1930 1959 1935 1944 1965 1946 1936 1939 1959 1952 1960 1958 1951 1963 1964 1938 1964 1943 1956 1953 1956 1942 1960 1957 1963 1938 1966 1961 1960 1964 1963 1960 LIST OF MEMBERS. 243 Douglas, Geoffrey William, B.Agr.Sc., c/- The Keith Turnbull Research Station, Private Bag, Frankston, Victoria. Durie, Peter Harold, M.Sc., C.S.1.R.O., Veterinary Parasitology Laboratory, Yeerongpilly, Brisbane, Queensland. Dyce, Alan Lindsay, B.Sc.Agr., 48 Queen’s Road, Asquith, N.S.W. Edwards, Dare William, B.Sc.Agr., Forestry Commission of N.S.W., Division of Wood Technology, 96 Harrington Street, Sydney. Endean, Robert, M.Sc., Ph.D., Department of Zoology, University of Queensland, St. Lucia, Brisbane, Queensland. English, Miss Kathleen Mary Isabel, B.Se., 6/168 Norton Street, Leichhardt, N.S.W. Evans, Miss Gretchen Pamela, M.Sc., Box 92, P.O., Canberra, A.C.T. Facer, Richard Andrew, ‘Moppity’, Parsonage Road, Castle Hill, N.S.W. *Fairey, Kenneth David, Box 1176, G.P.O., Sydney. Filewood, Lionel Winston Charles, c/- Department of Agriculture, Stock and Fisheries, Konedobu, Papua. Florence, Ross Garth, M.Sc.For., Ph.D., Forest Research Station, Beerwah, Queensland. Ford, Miss Judith Helen, 18 Central Avenue, Mosman, N.S.W. Fraser, Miss Lilian Ross, D.Se., ““Hopetoun”, 25 Bellamy Street, Pennant Hills, N.S.W. Gardner, Mervyn John, B.Sc.For., Dip.For., Forestry Office, Wauchope, N.S.W. *Garretty, Michael Duhan, D.Sc., Box 763, Melbourne, Victoria. Greenwood, William Frederick Neville, 11 Wentworth Avenue, Waitara, N.S.W. Griffin, David Michael, M.A., Ph.D. (Cantab.), School of Agriculture, Sydney University. *Criffiths, Mrs. Mabel, B.Sc. (née Crust), 50 Amourin Street, Brookvale, N.S.W. Griffiths, Mervyn Edward, D.Se., Wildlife Survey Section, C.S.1.R.0., P.O. Box 109, City, Canberra, A.C.T. *Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., D.T.M. & H. (England), 29 Flaumont Avenue, Lane Cove, N.S.W. Hadlington, Phillip Walter, B.Sc.Agr., 129 Condamine Street, Balgowlah, N.S.W. Hannon, Miss Nola Jean, B.Sc., Ph.D., 22 Leeder Avenue, Penshurst, N.S.W. Hartigan, Desmond John, B.Sc.Agr., 75 Northwood Road, Northwood, Lane Cove, N.S.W. Hennelly, John Patten Forde, B.Sc., Highs Road, West Pennant Hills, N.S.W. Hewiit, Bernard Robert, B.A. (Qld.), B.Sc. (Syd.), M.Se. (N.S.W.), A.R.A.C.I., Depart- ment of Chemistry, University of Malaya, Kuala Lumpur, Malaya. Hewson, Miss Helen Joan, B.Sc. (Hons.), School of Biological Sciences, Department of Botany, Sydney University. Higginson, Francis Ross, B.Sc.Agr. (Hon.), 3 Benson Street, West Ryde, N.S.W. Hill, Miss Dorothy, M.Sc, Ph.D., Department of Geology, University of Queensland, Brisbane, Queensland. Hindmarsh, Miss Gwenneth Jean, B.Sc., Botany Department, University College of Townsville, Pimlico, Townsville, Queensland. *Hindmarsh, Miss Mary Maclean,-B.Se., Ph.D., 4 Recreation Avenue, Roseville, N.S.W. *Holder, Miss Lynette Anne, B.Sc., 48 Rutiedge Street, Eastwood, N.S.W. *Hotchkiss, Professor Arland Tillotson, M.S., Ph.D. (Cornell), Department of Biology, University of Louisville, Louisville, Kentucky, 40208, U.S.A. *Hotchkiss, Mrs. Doreen Elizabeth, Ph.D., B.A., M.A., (née Maxwell), 2440 Longest Avenue, Louisville, Kentucky, 40208, U.S.A. Humphrey, George Frederick, M.Sc., Ph.D., C.S.I.R.O. Marine Biological Laboratory, Box 21, Cronulla, N.S.W. Ingram, Cyril Keith, B.A., B.He., 7 Ramsay Road, Pennant Hills, N.S.W. Jackes, Mrs. Betsy Rivers, B.Sc., Ph.D. (Univ. Chicago) (née Paterson), 114 Wellington Street, Aitkenvale, Hermit Park, North Queensland. Jacobs, Miss Janice Lorraine, B.Sc., School of Biological Sciences, Department of Botany, Sydney University. Jacobs, Maxwell Ralph, D.Ing., M.Sc., Dip.For., Forestry and Timber Bureau, Canberra, ALEC: Jacobs, Surrey Wilfrid Laurence, 7 Yarrara Road, Pymble, N.S.W. Jacobson, Miss Constance Mary, M.Sc., Ph.D., School of Biological Sciences, Department of Zoology, Sydney University. James, Sidney Herbert, M.Sc., 54 Holmfirth Street, Mt. Lawley, Western Australia. Jancey, Robert Christopher, M.Sc., Ph.D., P.O. Box 87, Rozelle, N.S.W. Jefferies, Mrs. Lesly Joan, 14 Denman Street, Hurstville, N.S.W. Jenkins, Thomas Benjamin Huw, Ph.D., Department of Geology and Geophysics, Sydney University. 244 1952 1947 1945 1960 1958 1963 1930 1960 1949 1951 1937 1966 1957 1956 1964 1932 1934 1936 1964 1964 1943 1966 1957 1958 1951 1948 1961 1922 1966 1931 1948 1966 1957 1953 1964 1966 1951 1932 1957 1954 LIST OF MEMBERS. Jessup, Rupert William, M.Sc., 6 Penno Parade North, Belair, South Australia. Johnson, Lawrence Alexander Sidney, B.Sc., c.o. National Herbarium, Royal Botanic Gardens, Sydney. Johnston, Arthur Nelson, B.Sc.Agr., 99 Newton Road, Strathfield, N.S.W. Jolly, Violet Hilary, M.Sc., Ph.D., Metropolitan Water, Sewerage and Drainage Board, 314 Pitt Street, Sydney. Jones, Edwin Llewelyn, B.A., P.O. Box 196, Leeton 6S, N.S.W. Jones, Leslie Patrick, Department of Animal Husbandry, Sydney University. Joplin, Miss Germaine Anne, B.A., Ph.D., D.Sc., Department of Geophysics, Australian National University, Canberra, A.C.T. Judd, Howard Kenniwell, Minnamurra Falls Forest Reserve, Box 14, P.O., Jamberoo, N.S.W. Keast, James Allen, M.Sc., M.A., Ph.D. (Harvard), Professor of Vertebrate Zoology, Queen’s University, Kingston, Ontario, Canada. Kerr, Harland Benson, B.Sce.Agr., Ph.D., Summer Institute of Linguistics, P.O. Ukarumpa, E.H.D., Territory of New Guinea. Kesteven, Geoffrey Leighton, D.Sc., c.o. Division of Fisheries and Oceanography C.S.1.R.O., P.O. Box 21, Cronulla, N.S.W. Khan, Abdul Ghaffar, M.Sc., 177 Commonwealth Street, Surry Hills, N.S.W. Kindred, Miss Berenice May, B.Sc., The Institute for Cancer Research, 7701 Burholme Avenue, Fox Chase, Philadelphia, Pa., 19111, U.S.A. Langdon, Raymond Forbes Newton, M.Agr.Sec., Ph.D., Department of Botany, University of Queensland, George Street, Brisbane, Queensland. Lanyon, Miss Joyce Winifred, B.Sc., Dip.Ed., 35 Gordon Street, Eastwood, N.S.W. Lawson, Albert Augustus, “Rego House’’, 23-25 Foster Street, Sydney. Lee, Mrs. Alma Theodora, M.Se. (née Melvaine), Manor Road, Hornsby, N.S.W. Lee, David Joseph, B.Sc., School of Public Health and Tropical Medicine, Sydney University. Levy, Miss Margery Olwyn, B.Sc., Dip.Ed., Katoomba High School, Martin Street, Katoomba, N.S.W. ; Littlejohn, Murray John, B.Sc., Ph.D. (W.A.), Department of Zoology, University of Melbourne, Parkville, N.2, Victoria. Lothian, Thomas Robert Noel, Botanic Garaens, Adelaide, South Australia. Lovedee, Miss Lois Jacqueline, B.Sc. (A.N.U.), 54 Shortland Street, East Gosford, N.S.W. Luig, Norbert Harold, Ph.D., c.o. Faculty of Agriculture, Sydney University. Lyne, Arthur Gordon, B.Sc., Ph.D., C.S.I.R.O., Ian Clunies Ross Animal Research Laboratory, P.O. Box 144, Parramatta, N.S.W. Macdonald, Colin Lewis, 7 Watford Close, North Epping, N.S.W. Macintosh, Professor Neil William George, M.B., B.S., Department of Anatomy, Sydney University. Mackay, Miss Margaret Muriel, B.Sc. (Hons.) (St. Andrews), M.Sc. (Syd.), M.I.Biol., J.P., Department of Botany, University College of Townsville, P.O. Box 499, Townsville, North Queensland. Mackerras, Ian Murray, M.B., Ch.M., B.Sc., C.S.I.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Mahmood, Ahmed, M.Sc., C/- Department of Botany, University of Karachi, Karachi-32, West Pakistan. *Mair, Herbert Knowles Charles, B.Se., Royal Botanic Gardens, Sydney. Marks, Miss Elizabeth Nesta, M.Sc., Ph.D., Department of Entomology, University of Queensland, Brisbane, Queensland. Martin, Angus Anderson, B.Sc. (Hons.) (Rand.), Department of Zoology, University of Melbourne, Parkville, N.2, Victoria. Martin, Anthony Richard Henry, M.A., Ph.D., School of Biological Sciences, Department of Botany, Sydney University. Martin, Mrs. Hilda Ruth Brownell, B.Sc. (née Simons), c/- Mrs. H. F. Simons, 43 Spencer ‘Road, Killara, N.S.W. Martin, Peter Marcus, M.Sc.Agr., Dip.Ed., School of Biological Sciences, Department of Botany, Sydney University. Mather, Mrs. Patricia (née Kott), M.Sc., Ph.D. (W.A. and Q’ld), Department of Zoology, University of Queensland, St. Lucia, Queensland. McAlpine, David Kendray, M.Sc., 12 St. Thomas Street, Bronte, N.S.W. McCulloch, Robert Nicholson, M.B.E., D.Sc.Agr., B.Sc., Cattle Tick Research Station, Wollongbar, N.S.W. *McCusker, Miss Alison, M.Se., Botany Department, University College, Box 9184, Dar es Salaam, Tanzania. McDonald, Miss Patricia M., B.Sc., Dip.Ed., 33 Holdsworth Street, Neutral Bay, N.S.W. 1956 1965 1948 1957 GAG 1962 1944 1947 1952 1949 1948 1947 1965 1963 1958 1960 1964 1955 1961 1958 1944 1955 1926 1965 1949 1947 1925 1922 1935 1920 1912 1948 1950 1961 1927 1950 1952 1954 1940 1966 1962 1957 1964 1922 1962 LIST OF MEMBERS. 245 McGarity, John William, M.Sc.Agr., Ph.D., Agronomy Department, School of Rural Science, University of New England, Armidale 5N, N.S.W. McGillivray, Donald John, B.Se.For. (Syd.), Dip.For (Canb.), P.O. Box 107, Castle Hill, N.S.W. McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), Service des Haux et Foréts, B.P. 285, Noumea, New Caledonia. McKenna, Nigel Reece, Department of Education, Konedobu, Papua. MeMillan, Bruce, M.B., B.S., D.T.M.& H. (Eng.), D.A.P.& E., F.R.E.S., School of Public Health and Tropical Medicine, Sydney University. McWilliam, Miss Paulette Sylvia, 4 Kingslangley Road, Greenwich, N.S.W. Mercer, Professor Frank Verdun, B.Sec., Ph.D. (Camb.), 24 Trafalgar Avenue, Roseville, N.S. W. Messmer, Mrs. Pearl Ray, 64 Treatts Road, Lindfield, N.S.W. *Meyer, George Rex, B.Sc., Dip.Ed., B.A., M.Ed., 91 Bowden Street, Ryde, N.S.W. *Miller, Allen Horace, B.Sc., Dip.Ed., 6 College Avenue, Armidale 5N, N.S.W. Millerd, Miss Alison Adéle, M.Sc., Ph.D., C.S.I.R.O., Division of Plant Industry, P.O. Box 109, City, Canberra, A.C.T. Millett, Mervyn Richard Oke, B.A., 18 Albion Road, Box Hill, Hil, Victoria. Milward, Norman Edward, B.Sc. (Hons.), M.Sc., Department of Zoology, University College of Townsville, Pimlico, Townsville, Queensland. Moore, Barry Philip, B.Sc., Ph.D., D.Phil., C.S.I.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Moore, Kenneth Milton, Cutrock Road, Lisarow, N.S.W. Moore, Raymond Milton, D.Sc.Agr., 94 Arthur Circle, Forrest, Canberra, A.C.T. Moors, Henry Theodore, B.Sc., Department of Geology and Mineralogy, University of Melbourne, Parkville, N.2, Victoria. Morgan, Mrs. Eva, M.Se., 4 Haberfield Road, Haberfield, N.S.W. Morrison, Gordon Cyril, 1/80 Queenscliff Road, Queenscliff, N.S.W. Moss, Francis John, 37 Avenue Road, Mosman, N.S.W. Moye, Daniel George, B.Sc., Dip.Ed., 6 Kaling Place, Cooma North, N.S.W. Muirhead, Warren Alexander, B.Sc.Agr., C.S.I.R.O., Irrigation Research Station, Private Mail Bag, Griffith, N.S.W. Mungomery, Reginald William, c/- Bureau of Sugar Experiment Stations, 99 Gregory Terrace, Brisbane, Queensland. Murray, David Ronald, B.Sc., 14 Consul Road, Brookvale, N.S.W. Murray, Professor Patrick Desmond Fitzgerald, M.A., D.Se., Department of Zoolcgy, University of New England, Armidale 5N, N.S.W. Nashar, Mrs. Beryl, B.Sc., Ph.D., Dip. Ed. (née Scott), P.O. Box 28, Mayfield, N.S.W. Newman, Ivor Vickery, M.Se., Ph.D., F.R.M.S., F.L.S., School of Biological Sciences, Department of Botany, Sydney University. Nicholson, Alexander John, C.B.B., D.Sc., F.R.E.S., C.S.I1.R.O., Box 109, City, Canberra. ALCL: *Noble, Norman Scott, D.Se.Agr., M.Sc., D.I.C., Unit 16, 25 Addison Road, Manly, N.S.W. Noble, Robert Jackson, B.Sc.Agr. Ph.D., 324 Middle Harbour Road, Lindfield, N.S.W. North, David Sutherland, 42 Chelmsford Avenue, Lindfield, N.S.W. O’Farrell, Professor Antony Frederick Louis, A.R.C.Sc., B.Sc., F.R.E.S., Department of Zoology, University of New England, Armidale 5N, N.S.W. O’Gower, Alan Kenneth, M.Sc., Ph.D., 20 Gaerloch Avenue, Bondi South, N.S.W. O’Malley, Miss Doreen Theresa, 100 Hargrave Street, Paddington, N.S.W. Osborn, Professor Theodore George Bentley, D.Sce., F.L.S., 34 Invergowrie Avenue, Highgate, South Australia. Oxenford, Reginald Augustus, B.Sc., 107 Alice Street, Grafton 3C, N.S.W. Packham, Gordon Howard, B.Se., Ph.D., Department of Geology, Sydney University. Parrott, Arthur Wilson, ‘“‘Lochiel’, Wakapuaka Road, Hira, R.D., Nelson, New Zealand. *Pasfield, Gordon, B.Sc.Agr., 20 Cooper Street, Strathfield, N.S.W. Patrick, John William, B.Sc.Ag. (Hons.), School of Biological Sciences, Department of Botany, Sydney University. Payne, William Herbert, A.S.T.C., A.M.I.H.Aust., M.A.P.E., 250 Picnic Point Road, Pienic Point, N.S.W. Peacock, William James, B.Sc., Ph.D., C.S.I.R.O., Division of Plant Industry, P.O. Box 109, Canberra City, A.C.T. Pedder, Alan Edwin Hardy, M.A. (Cantab.), Department of Geology, University of New England, Armidale 5N, N.S.W. Perkins, Frederick Athol, B.Sc.Agr., 98 Bellevue Terrace, Clayfield, Brisbane, Queensland. Philip, Graeme Maxwell, M.Sc. (Melb.), Ph.D. (Cantab.), F.G.S., Department of Geology, University of New England, Armidale, 5N, N.S.W. 246 1947 1935 1938 1960 1962 1929 1951 1952 1955 1957 1953 1957 1961 1964 1946 1958 1932 1964 1960 1932 1962 1919 1965 1950 1964 1948 1930 1947 1959 1955 1953 1943 1945 1937 1960 1932 1956 1965 1935 1962 1965 1952 LIST OF MEMBERS. Phillips, Miss Marie BHlizabeth, M.Sc., Ph.D., Parks and Gardens Section, Department of the Interior, Canberra, A.C.T. Pope, Miss Elizabeth Carington, M.Sc., C.M.Z.S., Australian Museum, College Street, Sydney. Pryor, Professor Lindsay Dixon, M.Se., Dip.For., Department of Botany, School of General Studies, Australian National University, Box 197, P.O., City, Canberra, A.C.T. Racek, Albrecht Adalbert, Dr.rer.nat. (Brno, Czechoslovakia), School of Biological Sciences, Department of Zoology, Sydney University. Rade, Janis, M.Sc., Flat 28a, 601 St. Kilda Road, Melbourne, S.C.38, Victoria. Raggatt, Sir Harold George, K.B.H., C.B.E., D.Sc., 60 Arthur Circle, Forrest, Canberra, A.C.T. Ralph, Professor Bernhard John Frederick, B.Sc., Ph.D. (Liverpool), A.A.C.I., School of Biological Sciences, University of New South Wales, P.O. Box i, Kensington, N.S.W. Ramsay, Mrs. Helen Patricia, B.Sc (née Lancaster), C/- Botany Department, University College of North Wales, Bangor, Caerns., United Kingdom. Reed, Mrs. Beryl Marlene, B.Sc. (née Cameron), 145 Duffy Street, Ainslie, A.C.T. Reed, Hric Michael, 145 Duffy Street, Ainslie, A.C.T. Reye, Eric James, M.B., B.S. (Univ. Qld.), Entomology Department, University of Queensland, St. Lucia, Brisbane, Queensland. Reynolds, Miss Judith Louise, c.o. Department of Entomology, University of Illinois, Urbana, Illinois, U.S.A. Richards, Miss Aola Mary, M.Sc. (Hons.), Ph.D. (N.Z.), School of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington, N.S.W. Richardson, Barry John, 12 Bowden Street, Parramatta North, N.S.W. Riek, Edgar Frederick, B.Sc., Division of Entomology, C.S.I.R.O., P.O. Box 109, City, Canberra, A.C.T. Rigby, John Francis, B.Sc., U.S. Geological Survey Coal Geology Laboratory, Orton Hall, Ohio State University, 155 S. Oval Drive, Columbus, Ohio, 43210, U.S.A. *Robertson, Rutherford Ness, F.R.S., B.Se., Ph.D., F.A.A., Professor of Botany, University of Adelaide, Adelaide, South Australia. Rothwell, Albert, D.P.A., 11 Bonnie View Street, Cronulla, N.S.W. Salkilld, Barry William, Dip.Soc.Stud. (Univ. Syd.), 53 Wareemba Street, Abbotsford, N.S.W. *Salter, Keith Eric Wellesley, B.Sc., School of Biological Sciences, Department of Zoology, Sydney University. Sands, Miss Valerie Elizabeth, M.Sc., Genetics Unit, Medical Research Council, P.O. Box 913, Dunedin, New Zealand. *Scammell, George Vance, B.Sc., 7 David Street, Clifton Gardens, N.S.W. Selkirk, David Robert, School of Biological Sciences, Botany Building, Sydney University. *Sharp, Kenneth Raeburn, B.Sc.. Eng. Geology, S.M.H.E.A., Cooma, 4S, N.S.W. Shaw, Miss Dianne Margaret, B.Sc., Faculty of Agriculture, Sydney University. Shaw, Miss Dorothy Edith, M.Sc.Agr., Ph.D., Department of Agriculture, Stock and Fisheries, Port Moresby, Papua-New Guinea. Sherrard, Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney. Shipp, Erik, Ph.D., 23 Princes Street, Turramurra, N.S.W. Simons, John Ronald, M.Sc., Ph.D., 242 Kissing Point Road, Turramurra, N.S.W. Slack-Smith, Richard J., 7 Kingston Street, Shenton Park, Western Australia. Smith, Eugene Thomas, 22 Talmage Street, Sunshine, Victoria. Smith-White, Spencer, D.Se.Agr., F.A.A., 51 Abingdon Road, Roseville, N.S.W. Southcott, Ronald Vernon, M.B., B.S., 13 Jasper Street, Hyde Park, South Australia. Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), No. 1 George Street, Tenterfield, N.S.W. Staff, Ian Allen, B.Sc., Dip.Ed., Department of Botany, Life Sciences, Southern Illinois University, Carbondale, Illinois, U.S.A. Stead, Mrs. Thistle Yolette, B.Sc. (née Harris), 14 Pacific Street, Watson’s Bay, N.S.W. Stephenson, Neville George, M.Sc. (N.Z.), Ph.D. (Lond.), School of Biological Sciences, Department of Zoology, Sydney University. Stephenson, Professor William, B.Sc. (Hons.), Ph.D. (Durham, Eng.), Diploma of Education, member Aust. Coll. Educ., Fellow Zoological Society, Department of Zoology, University of Queensland, St. Lucia, Brisbane, Queensland. Still, Professor Jack Leslie, B.Se., Ph.D., Department of Biochemistry, Sydney University. Strahan, Ronald, M.Sc., Department of Zoology, School of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington, N.S.W. Straughan, Mrs. Isdale Margaret, B.Sc. (Hons.), Department of Zoology, University of Queensland, St. Lucia, Brisbane, Queensland. Sullivan, George Emmerson, M.Sc. (N.Z.), Ph.D., Department of Histology and Embryology, Sydney University. 1955 1962 1962 1965 1940 1950 1950 1956 1960 1949 1944 1943 1946 1921 1965 1964 1952 1949 1917 1930 1940 1934 1961 1952 1909 1946 1947 OA! 1966 1936 1947 1927 1941 1964 1965 1963 1946 1964 1963 1926 1962 1960 1954 LIST OF MEMBERS. 247 Sutherland, James Alan, B.A., B.Sc.Agr., “Skye”, Galloway Street, Armidale, 5N, N.S.W. Sweeney, Anthony William, Malaria Institute, Public Health Department, Rabaul, Territory of Papua and New Guinea. Swinbourne, Robert Frederick George, 4 Leeds Avenue, Northfield, South Australia. Talbot, Frank Hamilton, M.Sc., Ph.D., Australian Museum, P.O. Box A285, Sydney South, N.S.W. Taylor, Keith Lind, B.Sc.Agr., C/- C.S.I.R.O., Division of Entomology, P.O. Box 109, Canberra City, A.C.T. Tchan, Yao-tseng, Dr., és Sciences (Paris), Microbiology Laboratory, Faculty of Agri- culture, Sydney University. Thompson, Mrs. Joy Gardiner, B.Sc.Agr. (née Garden), 21 Middle Head Road, Mosman, N.S.W. Thomson, James Miln, D.Sc. (W.A.), Department of Zoology, University of Queensland, St. Lucia, Brisbane, Queensland. Thorne, Alan Gordon, B.A., 82 Ben Boyd Road, Neutral Bay, N.S.W. Thorp, Mrs. Dorothy Aubourne, B.Se. (Lond.), Ph.D., ‘‘Sylvan Close’, Mt. Wilson, N.S.W Thorpe, Ellis William Ray, B.Sc., University of New England, Armidale 5N, N.S.W. Tindale, Miss Mary Douglas, D.Sc., 60 Spruson Street, Neutral Bay, N.S.W. Tipper, John Duncan, A.M.1.H.Aust., Box 2770, G.P.O., Sydney. *Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., c/- Australian Museum, P.O. Box A285, Sydney South, N.S.W. Tucker, Richard, B.V.Sc., Dr.Vet.M., Department of Veterinary Anatomy, University of Queensland, Mill Road, St. Lucia, Queensland. Tuma, Donald James, B.Se., c/- C.S.I.R.O., Division of Fisheries and Oceanography, P.O. Box 21, Cronulla, N.S.W. Valder, Peter George, B.Sc.Agr., Ph.D. (Camb.), School of Biological Sciences, Depart- ment of Botany, Sydney University. Vallance, Professor Thomas George, B.Sc., Ph.D., Department of Geology and Geophysics, Sydney University. Veitch, Robert, B.Sc, F.R.E.S., 24 Sefton Avenue, Clayfield, Brisbane, Queensland. Vickery, Miss Joyce Winifred, M.B.E., D.Sc., Royal Botanic Gardens, Sydney. Vincent, Professor James Matthew, D.Sc.Agr., Dip.Bact., Department of Microbiology, School of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington, N.S. W. Y *Voisey, Professor Alan Heywood, D.Sc., School of Earth Sciences, Macquarie University, 171-177 Epping Road, Eastwood, N.S.W. Walker, Donald, B.Sc., M.A., Ph.D., F.L.8., 18 Cobby Street, Campbell, Canberra, A.C.T. Walker, John, B.Se.Agr., Biological Branch, N.S.W. Department of Agriculture, Private Mail Bag No. 10, Rydalmere, N.S.W. Walkom, Arthur Bache, D.Se., 5/521 Pacific Highway, Killara, N.S.W. Wallace, Murray McCadam Hay, B.Sc., C.S.I.R.O., W.A. Regional Laboratory, Nedlands, Western Australia. Ward, Mrs. Judith, B.Se., c/- H.E.€., Gowrie Park, Tasmania. Wardlaw, Henry Sloane Halcro, D.Se., F.R.A.C.I., 71 McIntosh Street, Gordon, N.S.W. Wass, Robin Edgar, B.Sc. (Hons., Q’ld), Department of Geology and Geophysics, Sydney University. Waterhouse, Douglas Frew, D.Sc., C.S.I.R.O., Box 109, Canberra, A.C.T. *Waterhouse, John Teast, B.Sc., School of Biological Sciences, University of New South Wales, P.O. Box 1, Kensington, N.S.W. Waterhouse, Professor Walter Lawry, C.M.G., D.Se.Agr., M.C., D.I.C., F.L.S., F.A.A., 30 Chelmsford Avenue, Lindfield, N.S.W. Watson, Professor Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney University. Webb, Mrs. Marie Valma, B.Sc., 8 Homedale Crescent, South Hurstville, N.S.W. Webby, Barry Deane, Ph.D., M.Sc., Department of Geology and Geophysics, Sydney University. Webster, Miss Elsie May, 20 Tiley Street, Cammeray, N.S.W. Wharton, Ronald Harry, M.Sc., Ph.D., C.S.1I.R.O., 677 Fairfield Road, Yeerongpilly, Queensland. White, Andrew Hewlett, B.Sc. (Syd.), L. A. Cotton School of Geology, University of New England, Armidale, N.S.W. White, Mrs. Mary Elizabeth, M.Sc., 7 Ferry Street, Hunter’s Hill, N.S.W. *Whitley, Gilbert Percy, F.R.Z.S., Australian Museum, College Street, Sydney. Whitten, Maxwell John, Ph.D., C.S.I.R.O., Division of Entomology, P.O. Box 109, City, Canberra, A.C.T. Wildon, David Conrad, B.Sc.Agr., Box 108, Rozelle, N.S.W. Williams, John Beaumont, B.Sc., University of New England, Armidale, 5N, N.S.W. 248 1954 1960 1952 1950 1947 1965 1964 1964 1965 1949 LIST OF MEMBERS. Williams, Mrs. Mary Beth, B.Sc. (née Macdonald), 902D Rockvale Road, Armidale 5N. N.S.W. Williams, Neville John, B.Sc., c/- Mr. Brian Williams, Albury North High School, Albury, N.S.W. Williams, Owen Benson, M.Agr.Se. (Melbourne), c/- C.S.I.R.O., The Ian Clunies Ross Animal Research Laboratory, P.O. Box 144, Parramatta, N.S.W. Willis, Jack Lehane, M.Sce., A.A.C.I., 26 Inverallan Avenue, Pymble, N.S.W. Winkworth, Robert Ernest, P.O. Box 77, Alice Springs, Northern Territory, Australia. Woodward, Thomas Hmmanuel, M.Sc. (N.Z.), Ph.D. (Lond.), D.1.C., Department of Entomology, University of Queensland, St. Lucia, Brisbane, Queensland. Wright, Anthony James Taperell, B.Sc., Geology Department, Victoria University of Wellington, P.O. Box 196, Wellington, C.1, New Zealand. Yaldwyn, John Cameron, Ph.D. (N.Z.), M.Sc., Australian Museum, College Street, Sydney. Young, Graham Rhys, 8 Spark Street, Earlwood, N.S.W. CORRESPONDING MEMBER. Jensen, Hans Laurits, D.Sc.Agr. (Copenhagen), State Laboratory of Plant Culture. Department of Bacteriology, Lyngby, Denmark. 249 LIST OF PLATES PROCEEDINGS, 1966 I.—Fig. 1. Audiospectrograms of part of call of A, Hyla jervisiensis ; and B, H. ewingi; Fig. 2. Adult male Hyla jervisiensis. Ii.—Fenestella horologia and Minilya duplaris. IiI.—Cancrinella gyrandensis and Grantonia cracovensis. IV.—Anopheles atratipes Skusa and Anopheles tasmaniensis Dobrotworsky. V.—Anopheles tasmaniensis Dobrotworsky and Aedes nigrithorax Macquart. VI.—Hypopterygium rotulatum, meiosis in spore mother cells. 2,200. VII.—Hypopterygium rotulatum, meiosis in spore mother cells. VIII.— Hypopterygium rotulatum, leaves. TX.—Hypopterygium rotulatum, cells of leaves. LIST OF NEW GENUS, NEW SPECIES AND NEW SUBSPECIES Wo, Ga New Genus Page Novotettiz (Rhaphidophoridae) .. ane ra rs ie 5 ee LO) New Species Page Page cordicollis (Idacarabus) .. 179, 180 picta (Austroargathona) .. ey 16 cracovensis (Grantonia) .. TOS spelunecarius (Thenarotes) oF Till gyrandensis (Cancrinella) ee storyi (Acacia) .. ls 3 way leucoclada (Acacia) ai .. 149 subterraneus (Anomotarus) 4a 183 naracoortensis (Novotettiz) nao New Subspecies irrorata velutinella (Acacia) .. 147 leucoclada leucoclada (Acacia) .. 149 leucoclada argentifolia (Acacia).. 151 250 Page Abstract of Proceedings . 233 Acacia from eastern Australia, new taxa of, No. 2 147 Anderson, D. T., The comparative early embryology | of the Olgochaeta, Hirudinea and Onychophora (Presi- dential Address) : 10 Anderson, R. H., appear Honorary Secretary : 233 Annual General Meeting 1 Application of the names Sericesthis pruinosa (Dalman) and Sericesthis nigrolineata Boisduval ao Jey Atopozoa deerata (Sluiter): a discussion of the ae of the genus and species . 185 Australia, eastern, new taxa of Acacia from, No. 2 .. so day Australian cave Carabidae, new ao LY Australian freshwater prawns, a new corallanid isopod parasitic on 5 Lae Australian Tertiary echinoid genera, notes on three recently proposed .. 114 Baker, E. P., Inheritance of resistance to bunt in Turkey wheat selections.. 189 Baker, E. P., and Upadhyaya, Y. M., Studies on the inheritance of rust resistance in oats, IV 155 Balance Sheets for the year ending 28th February, 1966 7-9 Basden, R., The composition, occurrence and origin of lerp, the sugary secretion of Hurymela distincta (Signoret) : ae ae .. 44 Bass Strait Islands, Mosquitoes of Tasmania and ; 121 Baur, G. N., elected a member of ‘Clemeiil 236 Birds at Mitchell River Mission during the wet and dry seasons, rhino- nyssine nasal mite infestations in .. 211 Brachiopods, Permian, from Queensland, two new species of . 96 Breeding biology and larval development of Hyla jervisiensis .. 47 Britton, EH. B., The application of the names Sericesthis pruinosa (Dalman) and Sericesthis nigrolineata Boisduval 152 Browne, W. R., retirement from position of Honorary Secretary .. 233 Bunt in Turkey wheat selections, inheritance of resistance to oo 1K) Page Carabidae, new Australian cave. . .. 179 Carolin, R. C., Seeds and fruit of the Goodeniaceae, 58—see Notes and Exhibits. Catcheside, D. G., Sir William Macleay Memorial Lecture, 1966. The Centenary of Mendel 5 UO Chromosome numbers in Lamprothamnium 118 Comparative early embryology of the Oligochaeta, Hirudinea and Onychophora. . : 10 Composition, occurrence and origin of lerp, the sugary secretion of Hurymela distincta (Signoret) : 44 Conservation Photographic Exhibition... 2 Conversazione 2 Corallanid isopod parasitic on Australian freshwater prawns, a new . 176 Dandie, Alison K., appointed Linnean Macleay Fellow in Botany for 1967.. 238 Dobrotworsky, N. V., Mosquitoes of Tasmania and Bass Strait Islands .. 121 Domrow, R., Rhinonyssine nasal mite infestations in birds at Mitchell River Mission en the wet and Gh seasons 211 Echinoid genera, Australian Tertiary, notes on three recently proposed .. 114 Elections z ; 3, 233-234, 236-238 Exhibits—see N ores and Exhibits. Fauna Protection Panel, support given to the nominations of the two appointed by Chief Secretary to fill two vacancies on . 2 Fenestella horologia Bretnall and Minilya duplaris Crockford, on the species... 90 Fungi attacking seeds in dry seed-beds.. 84 Godwin, Professor H., Lecture on Radio - carbon Dating in the Quaternary .. oe a4 Goodeniaceae, seeds and Feuit of the . 58 Griffin, D. M., Fungi na ane seeds in dry seed-beds 84 Hansford, C. G., reference to death .. 234 Hewitt, B. R., ’ Highlands Peat of the Malayan Peninsula .. 231 INDEX 251 Page Page Hirudinea and Onychophora, the com- New taxa of Acacia from eastern parative early embryology of the Australia, No. 2 147 Oligochaeta 10 Notes and Exhibits : Hotchkiss, A. T., Chromosome numbers Carolin, R. C.—Film, prepared by in Lamprothamnium 118 himself and Mr. R. J. Oldfield, Hyla jervisiensis, the breeding biology demonstrating pollinating mech- and larval development of 47 anisms in Australian native plants 240 Hypopterygium rotulatum (Hedw.) Brid., Jacobs, 8. W. L.—Photographs intraspecific polyploidy in 220 ulustrating the distribution of starch in transverse sections of Inheritance of resistance to bunt in fresh grass leaves. 240 Turkey wheat selections 189 Khan, A. G.—Embryos of Podo- Intraspecific polyploidy in Hypopter ygium carpus species grown in agar rotulatum (Hedw.) Brid. .. 220 culture, 234—photographs and Isopod, new corallanid, parasitic on photomicrographs concerning the Australian freshwater prawns 176 nodulation of roots in three species of Podocarpus F 239 Jacobs, S. W. L., see Notes and Exhibits. Troughton, Ellis—Skull of the adult Johnson, L. A. S., elected a member of 3 holotype of the New Guinea Council : 1 Highland Dog, Canis hallstromi, from the MHuri-Duna country, Khan, A. G., see Notes and Exhibits. Southern Highlands district of Kosciusko State Park, appointment of Papua : 238 Dr. M. F. Day as a Trustee. 235 Vallance, T. G. —_His copy of George Kott, Patricia, Atopozoa deerata (Sluiter) ; : Shaw’s “Zoology of New Holland”’ a discussion of the relationships of (London, 1794) and a MS letter the genus and species 185 from Shaw to the artist James Sowerby giving instructions for Lamprothamnium, chromosome numbers an illustration in the book 239 in ; oa JING Whitley, G. P.—Note on announce- Lecturettes 5 1, 234, 236-238 ments of the formation of the Lerp, the sugary secretion of ‘Eurymela Linnean Society of New South distincta (Signoret), the composition, Wales in The Sydney Morning occurrence and origin of .. .. 44 Herald newspapers of October and Library accessions and report .. 2, 233-238 November, 1874 : 235 Linnaeus, portrait of, received as a gift Notes on three _ recently proposed from Linnean Society of London .. 2 Australian Tertiary echinoid genera 114 Linnean Macleay Fellowships : Report of work of Mr. A. J. T. Oats, studies on the inheritance of rust Wright, Fellow for 1965, 3—No resistance in, LV. . 155 appointment for 1966, 3—applica- Obituary Notices : tions invited for 1967, ce G. H. H. Hardy, 5; Miss Florence ment for 1967 ; 238 Sulman, M.B.E., 6;