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Proceedings of the 


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ket 
oo) 


Linnean Society 
of New South Wales 


ssued 17th June, 1971 


| Marine Biological Laboratory 
LIBRARY 
SEP { 71971 
WOODS HOLE, MASS. | 


VOLUME 96 
PART | 
No. 425 


Registered for posting as a periodical—Category A. 


The Linnean Society of New South Wales 


Founded 1874. Incorporated 1884 


‘‘ For the cultivation and study of the science of Natural History — 
in all its branches ”’ 


OFFICERS AND COUNCIL, 1971-72 


President 
L. A. 8S. Johnson, B.Sc. 


Vice-Presidents 


N. G. Stephenson, M.Se., Ph.D.; T. G. Vallance, B.Sc., Ph.D. ; 
and Miss Elizabeth C. Pope, M.Sc., C.M.Z.8., D. T. Anderson, D.Sc. 


Honorary Treasurer 
Joyce W. Vickery, M.B.E., D.Se., F.L.S. 


Secretary 
Mrs. Ruth J. Inall 


Couneil 
D. T. Anderson, D.Sc. E. Shipp, Ph.D. 
R. K. Bamber N. G. Stephenson, M.Sc., Ph.D. 
Barbara G. Briggs, Ph.D. F. H. Talbot, M.Se., Ph.D., F.L.S. 
W. R. Browne, D.Sc., F.A.A. EK. LeG. Troughton, C.M.Z.S., F.R.Z.S8. 
Rk. C. Carolin, Ph.D., B.Sc., A.R.C.S.  P. G. Valder, B.Sc.Agr., Ph.D. 
H. G. Cogger, M.Sc., Ph.D. T. G. Vallance, B.Sc., Ph.D. 
Mary M. Hindmarsh, B.Sc., Ph.D. Joyce W. Vickery, M.B.E., D.Sc., F.L.8. 
L. A. 8. Johnson, B.Sc. A. B. Walkom, D.Se. 
Elizabeth C. Pope, M.Sc., C.M.Z.S. G. P. Whitley, F.R.Z.S. 

Auditors 


W. Sinclair and Company, Chartered Accountants 


Iinnean Macleay Lecturer in Microbiology, University of Sydney 
K. Y. Cho, Ph.D. 


Innnean Macleay Fellow of the Society in Vertebrate Palaeontology 
as from Ist June, 1970 


Dr. A. Anne Warren (née Howie), B.Sc., Ph.D. (Cantab.) 


The Society’s Headquarters are in Science House, 157 Gloucester Street, Sydney, 
N.S.W. 2000, Australia 


ae 


Proceedings of the //.,.-\) 
Linnean Society uc” 
of New South Wales 


VOLUME 96 
No. 425-428 


CONTENTS OF PROCEEDINGS, VOLUME 96 


PART 1 (No. 425) 
(Issued 17th June 1971) 


CONTENTS 


Annual General Meeting : 
Report on the Affairs of the Society for the Year .. eh ne i 
Elections .. ee ae he ae te ge aie ae a 
Balance Sheets 5 


CAROLIN, R. C. The Trichomes of the Goodeniaceae  .. Si We 8 
CoGGER, H. G. The Reptiles of Lord Howe Island. (Plates I-l1) .. 23 
DARTNALL, A. J. Australian Sea Stars of the Genus Patiriella (Asteroidea, 
Asterinidae). (Plates t1-Iv) 1G Ae be i a barat oo) 
DARTNALL, A. J. A Sea Star of Genus Ctenodiscus from Tasmania deen a0) 


Hunt, G. 8S. The Genus Nunciella Roewer (Opiliones, Laniatores) with 
Description of a New Species from Kangaroo Island, South Australia 53 


PHiuip, G. M., and JAcKson, J. H. Late Devonian Conodonts from the 
Luton Formation, Northern New South Wales. (Plates yv-v1) i 66 


Erratum .. oe 2a ae ae aps Ai oe is ee 10 


Notes and instructions for authors .. ne ie ane a Ae 78 


PART 2 (No. 426) 
(Issued 15th September 1971) 


CONTENTS 


Bertus, A. L., and HAywarp, A. C. A Bacterial Leaf Spot of Zinnia 
in New South Wales. (Plates vul-vm1) 


ByRNES, N. A New Species of Lophopetalum (Celastraceae), a Genus 
New for Australia. (Plate Ix) 


MACLEAN, JOHN L. The Food and Feeding of Winter Whiting (Sillago 
maculata Quoy and Gaimard) in Moreton Bay 


TROUGHTON, ELLIs. The Early History and Relationships of the New 
Guinea Highland Dog (Canis hallstromt) 


WALKER, J. An Undescribed Species of Usiilago on Cyperus lucidus 
R.Br. in Australia. With comments on Robert Brown’s type 
collection of C. lucidus. (Plates X-xt1) 


WALKER, J., and Brertus, A. L. Shoot Blight of Hucalyptus spp. Caused 
by an Undescribed Species of Ramularia. (Plates x1-xm1) . 


Notes and instructions for authors 


Page 


81 


93 


99 


108 


AG, 


PART 3 (No. 427) 
(Issued 10th December 1971) 


CONTENTS 


BROADBENT, P., FRASER, L. R., and WATERWORTH, Y. The Reaction of 


Seedlings of Citrus spp. and Related Genera to Phytophthora 
citrophthora 


DEBENHAM, MARGARET L. Australasian Ceratopogonidae (Diptera, 
Nematocera). Part XV: The Genus Alluaudomyia Kieffer in 
Australia and New Guinea 


JACOBS, S. Systematic Position of the Genera Triodia R.Br. and 
Plectrachne Henr. (Gramineae) 


Lams, K. P. Aphids of Papua and New Guinea: An Annotated Check 
List 


WILLIAMS, O. B. Phenology of Species Common to Three Semi-arid 
Grasslands 


Page 


119 


128 


175 


186 


193 


PART 4 (No. 428) 
(Issued 17th April, 1972) 


CONTENTS 


CHIPPENDALE, G. M. Check List of Northern Territory Plants 


Howikz, ANNE. A Brachyopid Labyrinthodont from the Lower Trias 
of Queensland. (Plates XIV-XV) 


Martin, A. R.H. The Depositional Environment of the Organic Deposits 
on the Foreshore at North Deewhy, New South Wales 


O'REILLY, SUZANNE Y. Petrology and Stratigraphy of the Brayton 
District, New South Wales. (Communicated by Dr. R. E. Wass) 
(Plate XVI) 


TAYLOR, G. Sedimentation in Jervis Bay. (Communicated by Dr. 
K. A. W. Crook) 


Abstract of Proceedings 
List of Members 

List of Plates 

List of New Species 
Index 


Notes and instructions for authors 


Page 


207 


268 


278 


282 


297 
307 
313 
321 
321 
322 


331 


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ANNUAL GENERAL MEETING 
3lst MARCH 1971 


The Ninety-sixth Annual General Meeting was held in the Society’s Rooms, 
Science House, 157 Gloucester Street, Sydney, on Wednesday, 31st March 
1971, at 7.30 p.m. 

Professor N. G. Stephenson, President, occupied the chair. 

The minutes of the Ninety-fourth Annual General Meeting (25th March 
1970) were read and confirmed. 


REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 
1970 has seen a considerable number of changes and new developments 
in the Society’s affairs. So that members may be fully informed, this report 
will be more comprehensive than usual. 


Publication 

The Society’s Proceedings for 1969, Parts 2 and 3 were published on Ist 
June and 28th September, and for 1970, Parts 1 and 2 on 30th November and 
23rd December 1970. 

In order to retain registration with bulk postage facilities, the Proceedings 
will be published four times a year from 1971 onwards; during March, June, 
September and December. 

Donations totalling $1,000 were made during the year towards the cost of 
publication of three papers; net cost involved in publishing the Proceedings 
was $4,873.60. 


Membership 

During the year 11 new members were admitted to the Society, 1 died, 
10 resigned and 2 were removed from the list of members. The numerical 
strength of the Society at 1st March 1971 was: Ordinary Members, 272; Life 
Members, 30; Corresponding Member, 1; total, 303. 

The death of Dr. H. 8S. Halero Wardlaw in Singapore on 6th June 1970 is 
recorded with great regret. Dr. Wardlaw had been a member of the Society since 
1911, a member of Council from December 1924 until March 1970, and President 
during 1929/30. 


Administrative Changes 


At the beginning of the year Council decided to appoint a full-time Secretary 
to the Society. Mrs. Ruth J. Inall took up her appointment on 18th May 1970, 
combining the duties of administrator and editor of the Proceedings. 


Papers Read 


Papers read at Ordinary General Meetings totalled 18. Addresses were 
given at the following meetings: March, Early History of Australian Zoology 


PROCEEDINGS OF THE LINNEAN SocreTy oF NEw Sours Watezs, Vot. 96, Part 1 


2 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 


from earliest times up to and including Cook’s Voyages, by Mr. G. P. Whitley ; 
April, Carl Linnaeus, his personality, his relationships with contemporary 
botanists and students, his published works, his discovery of the sexual system 
of plant reproduction, and his introduction of the binomial system to scientific 
nomenclature by adapting the colloquial method of defining each material 
article by a general (generic) and a descriptive or qualifying (specific) word, by 
Dr. William T. Stearn; June, The biology of Nectaries, by Professor F. V. 
Mercer; September, The search for Australian fossil vertebrates, by Dr. A. 
Ritchie and Dr. Anne Howie; October, Cattle grazing and butterfly 
conservation—an exercise in conservation research in England, by Mr. O. B. 
Williams. Notes and exhibits were a feature of the November meeting. 


The Seventh Sir William Macleay Memorial Lecture was delivered by 
Sir Otto Frankel, D.Sc., F.R.S., F.A.A., on 27th July 1970, and was entitled 
“ Variation—the Essence of Life ”’. 


Archives 

The Society’s records have been sorted with the help of the Senior Archivist 
of the Archives Authority of New South Wales and a trained archivist from the 
Mitchell Library. The material is now housed, on permanent loan, at the Mitchell 
Library, where it will be catalogued, repaired and prepared for binding, and 
eventually available for reference to interested research workers. 


Science House and Future Developments Concerning the Society’s Activities 


The Sydney Cove Redevelopment Authority first informed this Society 
in June 1970 that it was intended to resume the property known as Science 
House at an early date; subsequently notice of resumption was received in 
November allowing 21 days for appeal against this action. 


During a cordial two-hour discussion at the North Sydney offices of 
the Redevelopment Authority, a number of matters relating to the possible 
future of the Society were discussed. It was necessary to ascertain whether the 
Sydney Cove Redevelopment Authority would give any encouragement to the 
Linnean Society or any other Learned Societies to remain in the eastern Rocks 
Area. In other words, we wished to determine whether or not the Authority, 
having threatened to destroy a Science Centre by resumption, would facilitate 
the redevelopment of a Science Centre in the area, either on the present or 
another site. 

In the meantime the Linnean Society had indeed been fortunate in that 
the Director of the Australian Museum, Dr. F. H. Talbot, had generously offered 
to provide, through the Australian Museum, accommodation for the Linnean 
Society and its Library in a new multi-storied block which is currently being 
planned as part of the Australian Museum complex of buildings. It is necessary 
for the Linnean Society to make a decision, without undue delay, as to whether 
it wishes to take advantage of this offer. 


Panbslitaais important to clarify the relationship between the Sydney Cove 
Redevelopment Authority and the Government, as well as the nature of the 
assistance which might be sought from either of these bodies. The Authority 
is an independent body, set up by the Government to develop a particular area. 
It hopes to do so under financially advantageous terms, but any profits made 
from the redevelopment are apparently to be returned to the Government by 
being paid into Consolidated Revenue. Thus, the Linnean Society could 
approach the Authority on matters concerning the resumption, or the provision 
of an alternative site, or for concessions of one type or another, but it would 
need to approach the Government directly for financial support if the changing 
circumstances necessitated grants or loans to maintain or develop the established 
activities of the Society. 


PROCEEDINGS OF THE LINNEAN SocrETy or New SoutH Watss, Vou. 96, Part 1 


REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 3 


The Chairman of the Authority was extremely sympathetic to the Linnean 
Society’s needs, and to the problems posed by the resumption order. These 
needs were elaborated by the Secretary and Honorary Treasurer of the Linnean 
Society, who were fully conversant with the details and were able to provide 
a solid factual backing to our case. 


The primary question raised was, whether or not the Sydney Cove 
Redevelopment Authority would provide, in the east Rocks Area under its control, 
a site for a scientific centre. The answer was that this question could be safely 
answered in the affirmative even now. ‘The project, of course, would be dependent 
on Government approval and support, but it could be phased so as to cause the 
Minimum inconvenience during change-over operations to the societies primarily 
concerned. One important point emerging from the discussion was that no 
land would be sold by the Authority after its resumption, but would be made 
available by the Authority under lease. No doubt the terms could be favourable 
if the cause for which the site would be developed was a worthy one and of public 
interest. 


On the assumption that a suitable site could be provided for a Science Centre, 
the question then arose as to the conditions under which it could be developed. 
The resumed land of the area is to be made available under lease from the 
Authority. Once a site is allocated, it seems that the occupant or occupants 
may make proposals for its development and execute an approved plan. 
Presumably there would be no objection to the provision in a multi-storied block 
of office space for sub-letting, comparable to the present tenancy arrangements 
in Science House. 


The next step in the proceedings is likely to be a direct discussion with the 
State Government. The Linnean Society, with the goodwill of the Sydney Cove 
Redevelopment Authority and the enthusiastic interest and support of other 
societies, may approach this period of negotiation with confidence. 


A survey was taken by a market research consultant to ascertain the support 
of other professional and scientific organizations. In view of the response, it 
was decided to hold a meeting of possible co-owners as soon as possible, with a 
view to establishing an organization to proceed with the planning for a Science 
Centre. 


The Secretary will also press for an early meeting with the Deputy Premier, 
Mr. Cutler. 


Inbrary 


Council, at its last meeting, discussed the implications of the last month’s 
developments, with particular reference to the Library. 


After considerable discussion, it was decided to retain the Library if at all 
possible, and that work on the Library proceed accordingly. It was agreed that 
a librarian would have to continue to be employed from now on (on a part-time 
basis). 

Library accessions from scientific institutions and societies on the exchange 
list from 1st March 1970 to 28th February 1971 amounted to 1,802. 


The total number of borrowings from the Library of books and periodicals 
by members and institutions for the year was 290. 


es Linnean Macleay Fellowship 
Dr. A. Anne Howie was appointed Linnean Macleay Fellow in Vertebrate 
Palaeontology for one year as from 1st June 1970. During the year Dr. Howie 
has been undertaking field trips to Rewan Station in Queensland which, among 
other material, yielded a skull of the first labyrinthodont found in that State. 


PROCEEDINGS OF THE LINNEAN Soctmety or New SoutH Wates, Vot. 96, Part 1 


4 REPORT ON THE AFFAIRS OF THE SOCIETY FOR THE YEAR 


Linnean Macleay Lectureship in Microbiology 

Dr. K. Y. Cho, Linnean Macleay Lecturer in Microbiology, University of 
Sydney, as from 1st January 1969, reported as follows on his work for the year 
to December 1970. Previous work on the extremely halophilic bacteria was 
accepted for publication in the Journal of General Microbiology (62: 267-270, 
1970). Purification and characterization of the exoprotease in these extreme 
halophiles are still in progress. The information obtained may be helpful in 
understanding the evolution of enzymes. Other projects under investigation 
are the use of spermine for preserving fine structure in bacteria and the effect. of 
vinea alkaloids on Azotobacter vinelandit. 


Annual Elections 

No nominations of other candidates having been received, the Chairman 
declared the following elections for the ensuing year to be duly made: 

President: L. A. 8S. Johnson, B.Se.- 

Members of Council: Barbara G. Briggs, Ph.D. ; H. G. Cogger, M.Sc., Ph.D. ; 
Elizabeth C. Pope, M.Sec., C.M.Z.S.; P. G. Valder, B.Se.Agr., Ph.D. (Camb.) ; 
T. G. Vallance, B.Sc., Ph.D.; G. P. Whitley, F.R.Z.A. 

Auditor: W. Sinclair & Co. 

The Chairman then installed Mr. L. A. 8. Johnson as President. 

A vote of thanks to the retiring President, Professor N. G. Stephenson, was 
moved from the Chair and carried by acclamation. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 96, Part 1 


5) 


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PROCEEDINGS OF THE LINNEAN Society OF New SoutH Wats, VoL. 


BALANCE SHEETS 


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PROCEEDINGS OF THE LINNEAN Society or New SourH Wates, Vou. 96, Part 1 


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PROCEEDINGS OF THE LINNEAN SociEtTy oF New SoutH Watss, Vou. 96, Part 1 


THE TRICHOMES OF THE GOODENIACEAE 


R. C. CAROLIN 
School of Biological Sciences, University of Sydney 


[Accepted for publication 9th December 1970] 


Synopsis 

A survey of trichomes in the Goodeniaceae is reported. It is concluded that there are seven 
distinct types of trichome: (1) stellate-simple; (2) uniseriate multicellular; (3) branched 
multicellular; (4) T-shaped; (5) glandular; (6) dendritic; (7) Brunonia-type. Types (1), 
(5) and (6) are subdivided into subtypes. Hach type is morphologically distinct and therefore 
has had a long, separate evolutionary history. ‘The hair types confirm previous suggestions in 
systematic rearrangement and again call into question the groupings made by the most recent 
monographer (Krause). 


INTRODUCTION 


The trichomes of the Goodeniaceae have not been examined in detail before. 
Krause (1912) makes passing references to them in his introductory paragraphs 
but gives few illustrations. He uses the indumentum type in his species descrip- 
tions and keys but only distinguishes between glandular, stellate and simple 
types, and in Dampiera provides some illustrations of different sorts of dendritic 
hairs. In some cases he refers to a species as being glabrous when, in fact, it 
does have numerous microscopic hairs (e.g. G. ovata). Colloza (1907, 1908) 
mentions the trichomes in a very few cases and illustrates only those of Brunonia. 
Hummel (1962) does not appear to mention the family. 


The use of trichomes in taxonomic studies is well established by a long line 
of investigators, particularly in the family Gramineae. In other families the 
taxonomist is usually content to describe the macroscopic appearance and 
texture of the indumentum. Uphof (1962) draws attention to a few cases in 
which more detailed investigations have been carried out. It is primarily because 
of their potential use in the taxonomy of the family that the present investigation 
has been undertaken. 


MATERIAL AND M@THODS 


Most of the observations have been carried out on herbarium material. 
Thin, surface sections of the dry material were cut by hand with a sharp razor 
blade ; the section bleached and soaked in a mixture of commercial bleaching 
agent and weak detergent. The sections were then mounted in Gurr’s water- 
mounting medium without staining. In the few cases where fresh material was 
available the soaking in the bleach was omitted. In addition, transverse sections 
of the leaves and corollas were prepared by embedding in paraffin wax in the 
usual manner. These two sets of preparations were further supplemented by 
observations on the whole specimen, scraping the hairs off the epidermis with a 
razor blade and mounting in Gurr’s water-mounting medium. 

Some of the sections were stained to distinguish the cuticle ; the stains used 
were ammoniacal fuchsin and Auromine. The former stains the cuticle red 
and also shows a brilliant fluorescence, whilst the latter is visible only with 
fluorescence microscopy. 


PROCEEDINGS OF THE LINNEAN Society or New SourH Wates, Vou. 96, Part 1 


R. C. CAROLIN 9 


OBSERVATIONS 


Coopernookia: Three basically different types of trichomes are found in this 
genus. Uniseriate multicellular hairs (Fig. 5) are almost restricted to the leaf 
axil. They consist of a basal cell with a prominent cuticle and a varying number 
of other cells with thin walls and a cuticle that is only visible using fluorescence 
microscopy. The so-called stellate hairs (Carolin, 1968) consist of a basal group 
of cells without outgrowths but with heavily cutinized surfaces, surmounted 
by a series of thick-walled, thin-cuticled cells with long, stiff, pointed outgrowths 
(Fig. 1, F). Furthermore, there are glandular hairs present and, whilst the two 
former hair-types are more or less uniform throughout the genus, these latter 
show some variation. In C. barbata the head of the hair is unicellular (Fig. 6). 
C. chisholmii also has this form of glandular hair but they are much less frequent. 
In C. strophiolata, C. polygalacea and C. georgei the head is four-celled and so 
flattened as to appear almost peltate (Fig. 3, 5). 


Fig. 1. Stellate and simple hairs. A. Goodenia ovata. B. Diaspasis filufolia. C. Goodenia 
pinnatifida. D. Goodenia  stelligera. E. Goodenia subintegra. FE. Coopernookia  polygalacea. 
G. Scaevola tomentosa. 


Velleia: Only simple unicellular hairs are found in this genus, except for some 
uniseriate multicellular hairs in the leaf axils. The former have thick cell walls 
which extend down into the basal part; the cuticle is very thin, and scarcely 
visible using normal staining and lighting. The only variation in the microscopic 
form of the hairs is in the degree to which the surrounding epidermal cells are 
raised up into a sheath around the base of the hair. This is fairly conspicuous 
in V. hispida (as in Fig. 1, c) but much less so in V. paradoxa and V. trinervis 
with V. rosea and V. daviesii showing intermediate conditions. Indeed, the size 
of the sheath is variable in the one species to some extent, although V. hispida 
jn particular shows a constantly well-developed sheath. 


Goodenia: This large genus displays a wide variety of trichomes ; most species 
have tufts, however minute, of uniseriate multicellular hairs in the leaf axils ; it 
is, however, the surface trichomes to which we should divert attention. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wates, Vou. 96, Part 1 


10 TRICHOMES OF THE GOODENIACEHKAE 


AS A 


Fig. 2. Uniseriate multicellular and T-shaped hairs. A. Goodenia incana. 
hederacea. C. Goodenia mueckeana. D. Goodenia primulacea. EK. Pentaptilon careyt. 
heterochila. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 96, Part 1 


B. Goodenia 
F. Goodenia 


R. C. CAROLIN tat) 


Fig. 3. 
D. Goodenia 
stephensonii. 


Glandular hairs. A. Calogyne sp. B. Goodenia decurrens. C. Goodenia ovata. 
concinna. EK. Coopernookia  polygalacea. F. Scaevola  coriacea. G. Goodenia 
H. Pentaptilon careyi. I. Goodenia barilletti. J. Leschenaultia hirsuta. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WateEs, Vou. 96, Part 1 


12 TRICHOMES OF THE GOODENIACHAE 


Despite the inadequacies of Krause’s (1912) treatment (see Carolin, 1959, 
1968), it seems best to use this as a basis for reporting these findings. 


The various forms of the trichomes are shown in Figs. 1, 2 and 3, and Table 1 
indicates their occurrence in the species examined. The stellate type is virtually 
the same, although generally shorter, as the stellate type in Coopernookia. 
Submerged stellate can be looked upon as the stellate type which has lost the 
basal column of more heavily cutinized cells, or as a cluster of simple trichomes. 
The simple and ensheathed simple trichomes have the same form as those found 
in Velleia. The one-sided simple hairs appear to be unique to Goodenia; these 
hairs are appressed and apparently ensheathed on one side, but this is obscured 
by an outgrowth of the trichome more or less overlapping and obscuring the 
basal sheath (Fig. 1, E). G. mueckeana appears to have a unique form of simple 
hair in that it is T-shaped (Fig. 2, c). 


Fig. 4. Dendritic hairs and branched multicellular hairs. A. Dampiera  coronata-. 
B. Verrauaxia reinwardti. C. Dampiera tomentosa. D. Dampiera stricta. EK. Dampiera purpurea. 


The glandular hairs can be divided into those with a single-celled head 
and those with a multicellular head (Fig. 3, A-E). In a few multicellular headed 
hairs the head is flattened to give the peltate hair (Fig. 3, c), but in others the 
the number of tiers of cells in the head is multiplied considerably and the head is 
oval or spherical in shape. This latter type may have a stalk which is uniseriate 
to the base (Fig. 3, A) or multicellular at the base (Fig. 3, 1). 


Calogyne: This genus falls into two fairly well defined groups on the basis of 
hair type, although at first they appear similar in that both have glandular and 
simple hairs. 


C. berardiana has simple hairs more or less ensheathed on one side only (as in 
Fig. 1, E) and glandular hairs with single-celled heads. 


C. holiziana and C. purpurascens have simple, practically sheathless cells and 
glandular hairs with multicellular heads (Fig. 6, v, 2). 


PROCEEDINGS OF THE LINNEAN SocitETY or NEw SoutH WALES, Vou. 96, Part 1 


R. C. CAROLIN 1133 


1.5 
14i 


iii 


Fig. 5. Hair types, see Table IIT. 


PROCEEDINGS OF THE LINNEAN Society or New SourH Wats. Vot. 96, Part 1 


14 TRICHOMES OF THE GOODENIACHAE 


Catosperma: The single species C. goodeniaceum has a few short simple hairs, 
particularly on the sepals, and rather more numerous cottony, multicellular 
hairs. 

Diaspasis: The single species D. filiformis has very thick-walled unsheathed 
simple hairs in which the cuticle is raised into numerous warty outgrowths ; 
the wall is conspicuously striated. 


v3 


Fig. 6. Hair types, see Table III. 


Scaevola: This genus shows rather fewer hair types than the other large genus 
Goodenia. In fact, however, another subtype is found which is almost a 
characteristic feature of the genus, i.e. the simple hair with an uneven, warty 
surface (Fig. 1, B). The occurrence of hair types in Scaevola and its supposed 
relative Diaspasis is shown in Table 2. 

It can be seen from this that stellate hairs are very infrequent, found only in 
S. tomentosa (Fig. 1, G) of the species examined. Simple, smooth hairs are also 
rare, being found only in S. mollis. The glandular and peltate types have 
apparently the same construction as those of Goodenia. 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH Watss, Vot. 96, Part 1 


T Bd “96 “TOA “SHIVA HEAOg MAGN 10 ALAIOOG NVANNI'T FHL FO SONTATIOON | 


Distribution of Hair Type in Goodenia and Related Genera 


TABLE 1 


Name 


Stellate Submerged 


1.1 and 
2 


Stellate 
1.3 


Simple 
1.41 


En- 
sheathed 
Simple 

1.5.1 


One- 
sided 
Simple 
1.6 


Multi- 
cellular 
HU 


T-shaped Glandular 


IV 


Vil 


Glandular 
with 
Multi- 
cellular 
Head 

V.2i 


Glandular 
with 
Multi- 
cellular 
Head and 
Multi- 
cellular 
Base 
V.2.11 


Peltate 
V3 


GOODENIA 

Sect. Monochila 
G. viscida R.Br. ee 
G. xanthotricha De Vriese 
G. watsonit F. Muell. 
G. sericostachys Gardner . . 


Ser. Racemosae 

?, quadrilocularis R.Br. 
7. eremophila B. Pritzel . 
G. stapfiana Krause 
7. ramellii F. Muell. 
G, decurrens R.Br. 
G, bellidifolia Sm. . 
G. stelligera R.Br. 
G. rostrivalvis Domin 
G. dimorpha Maiden et 

Betche RS 4 
7. geniculata R.Br. 4 
G. primulacea Schlechtd, . 
7. robusta (Benth.) Krause 
7. affinis De Vriese 
G. heterophylla Sm. 
G. hederacea Sm. .. 
@. rotundifolia R.Br. 


Ser. Suffruticosae 
7. ovata Sm. 
7, varia R.Br. Be 
7. mueckeana F. Muell. 
(. amplexans F. Muell. 


Ser. Coeruleae 
7, azurea F. Muell. 
G. incana R.Br... d 
*, horniana F. Muell. et 
Tate ap a 
G, scaevolina F. Muell. 
G. coerulea R.Br. .. 10 
G. trichophylla De Vriese ex 
Benth. ° 55 


Ser. Foliosae 
G. mitchellii Benth. 
Gi. heterochila F. Muell. 
7. hispida R.Br. 
7. grandiflora Sims 
(r. chambersii F. Muell. 
#, macmillanit F, Muell. . . 
7. armstrongiana De Vriese 


Ser. Pedicellosae 
G. cycloptera R.Br. ne 
G. pinnatifida Schlechtd. 
G. pusilliflora F. Muell. 
1. elongata Labill. 
- glauca F. Muell. b 
G. subintegra F, Muell. ex 
Black 2 Ap 
t. lunata Black 
7. heteromera F. Muell. 
G. pulchella Benth. 
7. concinna Benth. 
7. micrantha Christ. et Ost. 
7. cirrifica F. Muell. 
G. vilmorinae F, Muell, 


Seet. Amphichila 
G. paniculata Sm. 
G. gracilis R.Br, 
. pumilio R.Br. 


Incertae Sedis 
. tenuiloba F. Muell. 


ip} 


COOPERNOOKIA 

C. polygalacea (De Vriese) 
Carolin . a ite 

C. barbata (R.Br.) Carolin 

C. strophiolata (F. Muell.) 
Carolin .. m0 pe 


VELLEIA 

V. hispida Fitzg. .. 
V. paradoxa R.Br. 
V. trinervis Labill. 
V’. rosea Sp. Moore 
V. daviesi: F. Muell. 


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Pentaptilon: This monospecific genus has very much the same indumentum as 
Verreauxia, all the same types of hair are present, simple, unbranched multi- 
cellular, branched multicellular, glandular with multicellular heads. 


VI.2 


== 
os 


Fig. 7. Hair types, see Table III. 


Leschenaultia: Few species of this genus appear to have trichomes except on 
the inner floral parts. JZ. hirsuta has glandular hairs with flattened multicellular 
heads on multicellular stalks. The heads are frequently knocked off and 


PROCEEDINGS OF THE LINNEAN SocireTy oF New Soutsa Watss, Vou. 96, Part 1 


if 


o2) 


TRICHOMES OF THE GOODENIACEAE 


consequently trichomes on the older parts of the plant often appear to be simple 
(Fig. 3, J). 

Dampiera: This large genus appears to show only the dendritic hair-types 
These hairs at first sight appear to be similar to those of Verreauxia but differ 
in that they have two short arms instead of one to each cell, except the basal 
two or three and the terminal cell. There is some variation on the basic theme 
of this hair-type in Dampiera which will no doubt prove of taxonomic value in 
the future. The variation is concerned mainly with the position of the two 
branches of each cell with regard to each other, the length of the cells and the 
number of the cells in the trichome. 


In D. discolor the arms are situated opposite each other and more or less 
in the middle of the cell (Fig. 7, v1.1), whilst in D. hederacea, D. incana and 
D. luteiflora they are situated at the top of the cell opposite each other (Figs. 4, 5, 
7, v1.2). In many species there is a tendency for the upper cells to be attached 
to the hair somewhat laterally rather than terminally on the preceding cell of 
the trichome, e.g. D. spicigera (Fig. 4, Cc). In D. coronata, D. lindleyi and D. 
eriantha there are only one to three armed cells, and the body of each cell is very 
short (Figs. 4, A; 7, v1.4). There is a tendency also for each cell to be displaced 
laterally, as in D. stricta (which see) but this is usually very slight. 

D. eriocephala and D. adpressa show the condition when the arms of each 
armed cell are not opposite but one is placed at the top of the cell and one at the 
bottom (Figs. 4,cC; 7, VI.3). 

In D. stricta the arms are also displaced but, in addition, each succeeding 
cell of the trichome is bent horizontally at its base, thus giving a series of steps 
(ame, 4, Ds Ws \it8))), 

Brunonia: This genus has, at first sight, a hair-type entirely different from 
that found in any other genus in the family. 

The hairs are multicellular and closely appressed to the epidermis. The 
basal cell is thin-walled and quite obviously cutinized, whilst all the succeeding 
cells are very thick-walled with a very thin cuticle. The first of these latter cells 
turns the trichome through 90° since it is attached horizontally to the basal cell 
(Fig. 7, vit). Moreover, it has a prominent groove in the upper surface, particu- 
larly towards the base. The next cell of the trichome is attached diagonally 
to the apex of the first and frequently the two spiral around each other for some 
distance (Fig. 7, vit). There are two subtypes, both found on the same plants. 
In one this second cell is the terminating cell and has large warty projections on 
the surface. In the second subtype, which is much longer, there are usually 
still more cells in the file, all attached to each other in much the same manner as 
indicated above, and none of them has prominent warty projections. 


DISCUSSION 
On the basis of these results it is possible to distinguish seven main trichome 
types : 
(i) Stellate-simple type in which one or several unbranched cells are 
arranged side by side in short multicellular files (Fig. 5, 1.1-1.5). 
(ii) Uniseriate multicellular type (Fig. 5, 11). 
(iii) Branched multicellular type in which each cell, except the basal and 
terminal ones, branch once (Fig. 7, 11). 
(iv) T-shaped type (Fig. 5, Iv). 
(v) Glandular type (Fig. 6). 
(vi) Dendritic type in which each cell except some basal ones and sometimes 
the terminal one has two side-arms (Fig. 7, v1.1—v1.5). 
(vil) Brunonia-type (Fig. 7, vit), which may be a derivative of (ii). 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vot. 96, Part 1 


R. C. CAROLIN 19 


Each of these trichome types appears to have had its own independent 
phylogenetic history within the family ; independent that is, as far as any part 
of the organism can be independent of any other part. There is, moreover, 
probably no significant phylogenetic link between the main types, i.e., we cannot 
look upon any of the seven types outlined above as being derived from each other, 
with the possible exception of the Brunonia-type. They all differ from each 
other fairly significantly. For instance, it is improbable that the thick-walled 
stellate-simple trichome of horizontally arranged cells gave rise to the single-file 
multicellular hair or to the thin-walled secretory glandular hairs. 


The dendritic type does show some similarity to the unbranched multi- 
cellular type in that they both consist of uniseriate files of cells with a basal group 
of heavily cutinized cells. It is felt, however, that any such derivation must 
have occurred at a very early stage in the history of the family since their 
appearances are now so different. There is a slight resemblance to the branched 
multicellular type, but this would appear to be rather superficial since there are 
no basal cells more heavily cutinized than the upper ones ; in fact, the insertion 
of these basal cells on the epidermis is more like that of simple hairs. 


The most complex of the stellate-simple type is found in Coopernookia 
(Fig. 5,1.1). It has been indicated previously that this genus has a number of 
features that may be considered primitive in the genus (Carolin, 1966). This is 
really the only evidence that is available for suggesting a phylogenetic starting 
point for the various forms of this trichome type. It is easy enough to arrange 
these types in a series (Fig. 5), but to determine whether it is a reduction series 
or compounding series other evidence is necessary. For the reason given it will 
be considered as a reduction series in which first the individual units are con- 
tracted into a submerged, unilayered, stellate form (1.2) then submerged in the 
epidermis (1.3) and then reduced to a single unit forming a simple trichome 
(1.4). Type 1.5 is similar but is ensheathed at the base by a raised plinth of 
cells. 

The appressed hairs of Goodenia glauca and its allies are derived from the 
simple form by the excessive growth of the plinth cells on one side (1.6). The 
warty type might almost be considered a subform of the simple trichome. In 
many ways the T-shaped trichomes of G. mueckeana (Fig. 5, Iv) could be mistaken 
for suppressed dendritic trichomes. However, it does not have the full comple- 
ment of basal cells for the dendritic type. Furthermore, this latter type is 
completely unknown in Goodenia, to which G. mueckeana undoubtedly belongs, 
or any of its close relatives. If, in fact, this trichome type is genetically related 
to the dendritic type, it is either a phylogenetic ‘ throw-back”’ of some 
significance or a feature which has been retained from the ancestral Goodeniaceae 
in only this one species over a very considerable period of time. In the absence 
of any further evidence probably the best course is to recognize it as a separate 
type. The presence of a basal cutinized cell may link this cell type to the 
unbranched multicellular type. 


The unbranched multicellular type shows relatively little variation in form 
throughout those genera in which it occurs. The variation that does occur is 
confined to differences in length and straightness of the hair, features possibly 
significant taxonomically but not particularly so in a general survey of the type 
being reported here. 


The dendritic type shows almost as much formal variation as the stellate- 
simple type. Again, it is almost impossible to decide which form is primitive ; 
possibly the equal-armed type with the arms at the mid-point of the cell could 
be taken as a starting point (Fig. 7, v1.1). Changes have been firstly the arms 
occurring at the top of each cell (Fig. 7, v1.2) and secondly one arm occurring 
at the top and one at the bottom (Fig. 7, v1.3). The suppression of the lower 


PROCEEDINGS OF THE LINNEAN SoctETy oF NEw SoutH Watss, Vou. 96, Part 1 


20 TRICHOMES OF THE GOODENIACEAE 


part of each cell and the extension of the arms leads to Fig. 7, v1.5, whilst v1.5 
has lead to form v1.6 by the whole trichome becoming appressed. 


The Brunonia type could have been derived from this last form, i.e., 6, by 
suppression of both arms or their connation, or from type u. The evidence 
available does not allow a decision to be made. 


Again, the glandular hair-type shows very considerable variation, and whilst 
it is easy enough to draw a series, the designation of the primitive condition is 
not so simple. Since the stalk is a file of cells, it seems reasonable to conclude 
that in the primitive condition the head will continue this sequence and also be 
unicellular. Thus this series can be looked upon as a compounding series, the 
head increasing in complexity. The first step is a double division of the terminal 
cell to give the situation in 2. In one line of change this becomes flattened into 
the peltate form of 3 and in another the multicellular form of 4 (all Fig. 6). The 
subforms of v.2 are based on the single or multicellular base. The form found 
in Leschenaultia is sub-peltate but with a long stalk. 


Any sort of overall concept of evolution within the family must await more 
complete evidence. This survey of trichome form and phylogeny is an attempt 
to determine the homologies of the various trichomes, a task which has to be 
performed before any overall taxonomic or phylogenetic assessment can be 
made. 


Functionally, it is possible that all these trichomes have a bearing on the 
adaption of these plants to the hot, dry, sunny climates in which many of them 
live. The non-glandular trichomes reflect incident light and heat and thus may 
restrict water loss. Various species have developed the appressed habit of 
trichomes, but using different types, thus in Goodenia glauca they are simple, 
in Goodenia hederacea they are uniseriate multicellular, in Dampiera stricta and 
D. coronata they are still another type ; the same apparent condition is achieved 
by non-homologous structures in these cases. 


The glandular hairs may likewise function in decreasing water loss since they 
secrete a resin which covers the epidermis and frequently even obstructs the 
stomata. 


More information is obviously necessary with regard to the physiology of 
these plants, but it is reasonable to assume that the trichomes do, in fact, have a 
Significance in the adaptation of these organisms. 


It seems clear that these trichomes will be important taxonomic characters. 
The last monograph of the family (Krause, 1912) made relatively little use of 
them. 


In some ways this investigation indicates that adjustments may be needed 
in previous suggestions (Carolin, 1959, 1966, 1967; Peacock, 1963), although 
they promise to be relatively minor. 


Verreauxia and Pentaptilon, otherwise clearly agreeing with Goodenia and 
its satellite genera, stand apart in their possession of branched multicellular 
hairs. Leschenaultia, likewise, shows an aberrant trichome type resembling 
those of the Goodenia group more than those of the Dampiera group. 

On the other hand the trichome types of Coopernookia, Goodenia, Velleia, 
Scaevola, Diaspasis, Catosperma and Calogyne are all very similar and, apart from 
the presence of branched multicellular types, those of Verreauxia and Pentaptilon 
are also similar to those of Goodenia. This 7- and 8-chromosome group (Peacock, 
1963), with ‘‘ polytelic’’ inflorescences (Carolin, 1967), ‘‘ Goodenioid ’”’ floral 
morphology (Carolin, 1959) and with a single ovular strand and frequently winged 
seeds (Carolin, 1966) has distinct similarities when the trichomes are considered. 
Diaspasis is clearly very close to Scaevola in its trichome type. 

The other genera, in the 9-chromosome group (Peacock, 1963), are less 
similar, as indeed has been shown in other previous studies (Carolin, 1959, 1966.) 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Watzs, Vou. 96, Part 1 


R. C. CAROLIN 21 


The trichomes of Brunonia may show certain similarities with those of Dampiera 
but they are not close (see above). 


The trichome type also promises to be of some use in the subdivision of the 
larger genera, particularly in Goodenia, and less so in Dampiera and Scaevola. 
A glance at Table 1 will show the occurrence of multicellular hairs to be concen- 
trated in the ser. Rosulatae and ser. Racemosae and peltate hairs occur in G. ovata 
and its relatives G. vernicosa and G. varia. The one-sided simple type likewise is 
concentrated in G. glauca and its relatives, whilst the other forms tend to be 
scattered. 


TABLE 3 
Refer to Figures. Standardized Hair Types in Goodeniaceae 


Description Type Form Subform Figure 

Stellate : I 1 1 5 
Reduced stellate I 2 1 5 
Submerged stellate : F I 3 1 5 
Submerged stellate, warty suns I 3 i 
Simple : if 4 1 5 
Simple, warty surface I 4 i 5 
Ensheathed simple .. : I 5 1 5 
Ensheathed simple, warty surface I 5 ul 
One-sided simple : i 6 1 5 
Multi-cellular unbranched ae §: II — - = 5 
Multi-cellular branched .. ae ah iil — — 7 
T-shaped ap Ne os oft re NVA = = 5 
Glandular... V 1 —_— 6 
Glandular with multi-cellular head : WW 2 1 6 
Glandular with multi-cellular head and 

roulti-cellular base Vv 2 u 6 
Peltate Vv 3 — 6 
Leschenaultia type d a W 4 — 6 
Dendritic, branched at said: cell VI 1 — 7 
Dendritic, branched at top of cell VI 2 — 7 
Dendritic, one branch at top and one 

bottom of cell .. Vi 3 — 7 
Dendritic with very short cells and long 

arms at = VI 4 — 7 
Dendritie with unequal arms te Bc WAL 5 — 7 
Brunonia-type 3a aa ay ae Vil — — 7 


It is clear once again that Krause’s subgeneric divisions have as little bearing 
on trichome distributions as they have on seed type and inflorescence type 
(Carolin, 1959, 1966). Little more can be said until a monograph is completed. 


In Scaevola there are few distinctions, but peltate forms occur in a fairly 
compact set of species grouped around S. porocarya. 


In Dampiera the only striking grouping of similar trichomes is type Iv.5 
in Sect. Camptosperma. 


ACKNOWLEDGEMENTS 


To Miss M. Burton for making many of the slides and completing some of 
the illustrations. Material used in this investigation was collected with the aid 
of A.R.G.C. and University of Sydney Research Grants. 


PROCEEDINGS OF THE LINNEAN Socrety oF New SoutH Wa tes, Vou. 96, Part 1 


22 TRICHOMES OF THE GOODENIACHAE 


Bibliography 
Caron, R. C., 1959.—F loral Structure and Anatomy in the family Goodeniaceae. Proc. Linn. 
Soc. N.S.W., 84: 282. 
, 1967. The Concept of the Inflorescence in the Order Campanulales. JIbid., 92: 7. 
, 1966. Seeds and Fruits of the Goodeniaceae. Jbid., 91: 58. 
Cottoza, A., 1907, 1908.—Studio anatomico sulle Goodeniaceae, Nuovo. Giorn. Bot. Ital., 14: 
304, 15: 5. 
Hummet, K., 1962. Die Verbreitung der Haartypen in den Naturlichen Verwandtschafts- 
gruppen. Hncycl. Pflan Anat., Berlin. 
Krause, K.. 1912. Goodeniaceae und Brunoniaceae. Das Pflanzenreich, 54, Berlin. 
Peacock, W. J., 1963.—Chromosome numbers and cyto-evolution in the Goodeniaceae. Proc. 
Lunn. Soc. N.S.W., 88: 8. 
Upnor, J. C. Th., 1962.—Plant Hairs. Hncycl. Pflan. Anat., Berlin. 


PROCEEDINGS OF THE LINNEAN Society or NEw South WaAtEsS, VOL. 96, Part 1 


THE REPTILES OF LORD HOWE ISLAND 


H. G. COGGER 
Australian Museum, Sydney 


(Plates 1, 1) 


[Accepted for publication 9th December 1970] 


Synopsis 

Three species of lizards are the only terrestrial reptiles recorded from Lord Howe Island. 
Of these, only the gecko Phyllodactylus guenthert Boulenger and the endemic skink Leiolopisma 
lichenigera (O’Shaughnessy), both of which are redescribed, are currently known to occur on the 
island. 

Although both lizards are abundant on the small islands lying off-shore from Lord Howe 
Island (including Ball’s Pyramid, from which both species are here recorded for the first time), 
they are now uncommon on the main island and confined to a small specialized habitat. The 
latter is described. 

Records of the gecko Gehyra oceanica (Lesson) from Lord Howe Island are shown to be in 
doubt. 


INTRODUCTION 

Lord Howe Island (Fig. 1, Pl. 1, A, B) is a crescent-shaped island roughly 
74 miles long and averaging about one mile in width, and is located approximately 
440 miles east-north-east of Sydney, New South Wales. Its highest point 
(Mount Gower) rises to 2,836 feet above sea-level. The island is the central 
portion of a platform which has an average depth of about 50 fathoms. Standard 
(1961) considered that this platform was formed by wave action during the 
Pleistocene. 

Lord Howe Island has long held great interest for the biologist, largely 
because of its isolation and the relatively high order of endemicity of its plants 
and animals. Until its discovery by Lt. Henry Lidgbird Ball in 1788 (only two 
weeks after the founding of the first Australian settlement at Port Jackson), 
Lord Howe Island had not known human occupation. 

The history of the island has been well documented by Nicholls (1938) and 
Hindwood (1940). Oliver (1917) gives an excellent account of the plants of the 
island, while Standard (1963) gives a full account of its geology. Standard 
(1961) discusses Lord Howe Island in relation to the submarine geology of the 
Tasman Sea. Excellent general accounts of the island and its natural history 
are given by McCulloch (1921), Pope (1949, 1959, 1960) and Paramonov (1958, 
1960 and 1963). The latter discussed briefly the status and habits of the island’s 
reptiles. More recent treatment of the birds is given by Hindwood and 
Cunningham (1950) and McKean and Hindwood (1965). Squires (1963) provided 
Carbon-14 datings of 20,700 years and more than 38,500 years for two samples 
from the Fossil Dune Sequence on Lord Howe Island, from which specimens 
of the large extinct turtle Meiolania platyceps have been taken. 

Significant changes have been wrought in the original environment of Lord 
Howe Island since European occupation. Eight species of land birds have 
become extinct (Hindwood, 1940 ; McKean and Hindwood, 1965). The endemic 
phasmid Dryococelus australis was also believed to be probably extinct (Gurney, 
1947), but this insect was recently rediscovered on Ball’s Pyramid, south of Lord 
Howe Island (McAlpine, 1966). There seems little doubt that this insect is 
extinct on the main island. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WatEs, Vou. 96, Part 1 


24 REPTILES OF LORD HOWE ISLAND 


The major disruption to the ecology of the island is thought to have been 
brought about by the chance introduction of the Black Rat (Rattus ratius) in 
1918. The Burns Philp steamship Makambo ran aground on 14th June of that 
year and rats apparently accompanied the loads of fruit and copra which were 


North Islet (7) 


ADMIRALTY 
ISLANDS 


278 
Roach |. 
Noddy |. & 31°30" 


® south. 
Sugarloaf |. ) 


Tenth of June |. 


® Soldier's Cop |. 


Mt. Eliza OAS SoeSsau (714") 
oo Gc ®@ 


Ned's Beach 


Dawson Ridge 


4 Phillip Rock Steven's Pt. 


Middle Beach 


Blinkeotoreesboy 2 Mutton Bird Island 


Mutton Bird Pt. 


Rocky Pt. 


Boat Haven 


Wolfe Rock 


KSalmon Beac'! 
SF Sa, East Pt. 
oo 


@ sr. Lidgbird (2504') 


Sugarloaf Pt. 


Red Pt. 


Erskine Valley > 


31°35! 


Et Gower (2835") 


Gower |. & 
159°05' 


Fig. 1. Map of Lord Howe Island showing topography and principal localities. Contour 
lines are at 500 ft. intervals. 


thrown overboard and washed ashore. At least, it was immediately after this 
event that the islanders first noticed the presence of rats. Within a short time 
the numbers of these animals had reached plague proportions and were having 
a disastrous effect on the island’s principal industry, the harvesting and export 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 96, Part 1 


H. G. COGGER 25 


of the seeds of the endemic palms (Howea spp.). For many years a bounty was 
offered on rats, but at the time of writing the number of rats is maintained at a 
relatively low level due to systematic poisoning by the Island Board. 


Prior to the introduction of the Black Rat, goats and pigs had been released 
and these were soon running wild (and still do) in the uninhabited parts of the 
island. 


Hindwood (1940) records that only three species of land birds were extinct 
when rats were first introduced in 1918, but by 1935 an additional five species 
had vanished. The extent to which rats were responsible for eliminating these 
birds is difficult to assess, but there seems little doubt that on the main island 
rats exterminated the endemic phasmid mentioned above. 


One of the aims of the author’s visit to Lord Howe Island was to determine 
whether the reptile fauna had been influenced significantly by any of those 
factors which had brought about changes in other animal groups. There was, in 
fact, some evidence to suggest that changes had occurred in the reptile population, 
for recent reports from the island suggested that lizards were only infrequently 
seen and difficult to collect on the main island, despite the fact that they were 
extremely common on Rabbit Island (Pl. 1, c). The latter is a low island, about 
250 yards long and 118 feet high, lying within the lagoon less than half a mile 
off-shore. 


( fo) 


MEAN TEMPERATURE 


4 5 6 7 


MEAN PRECIPITATION (inches) 


Fig. 2. Climatograph for Lord Howe Island based on data supplied by the Commonwealth 
Bureau of Meteorology. Months are indicated by numbers. 


No attempt has been made to treat the marine reptiles of Lord Howe Island. 
The sea-snake fauna of north-eastern Australia is a rich one, and although many 
species probably occur in the waters near Lord Howe Island, I can find no reliable 
records. Indeed, such records would probably bear little relationship to the 
number of forms occurring in the area. 


Similarly, all of the marine turtles known from the Great Barrier Reef 
probably frequent the waters of Lord Howe Island from time to time, but no 


PROCEEDINGS OF THE LINNEAN SoctmtTy or New SoutH Watss, Vou. 96, Part 1 


26 REPTILES OF LORD HOWE ISLAND 


reliable records are available. However, it is interesting to note that turtles, 
presumably the Green Turtle (Chelonia mydas), once nested on Lord Howe Island 
and provided a valuable source of food for the early visitors and colonists. 
Phillip (1789) notes that turtles were nesting in abundance in March 1788. 


Turtles no longer breed on the island and they are only infrequently seen 
in the surrounding seas. The disappearance of the turtle colony is generally 
attributed to excessive hunting, but this explanation is not entirely satisfactory. 
Turtles have been hunted intensively at their breeding sites for many years in 
many places throughout the world, and although serious depletion of breeding 
populations has often resulted, the total extermination of such populations 
has not proved to be readily accomplished. However, it may be that the isolation 
of Lord Howe Island prevented recolonization after its relatively small breeding 
community was reduced to a level which was no longer viable. 


Situated some 10 nautical miles to the south of Lord Howe Island is Ball’s 
Pyramid, a large eroded remnant of volcanic rock (of the same series as the 
Mt. Lidgbird volcanics of Standard, 1963) which rises to a height of 1,811 feet. 
This rock is still virtually unexplored biologically and was first scaled in 1965 by 
a party from Sydney Rock Climbing Club. Two earlier attempts by the same 
club were unsuccessful, and although no biological collections were made, on the 
second of these trips photographic records of the ‘‘ extinct ’’ Lord Howe Island 
phasmid Dryococelus australis (McAlpine, 1966) and the scincid lizard Letolopisma 
lichenigera (see later discussion) were made. Another climbing party in February 
1969 collected specimens of both Letolopisma lichenigera and Phyllodactylus 
guentheri. 

Ball’s Pyramid sits on a shallow platform which is separated from that of 
Lord Howe Island by depths of more than 2,000 feet and was probably never 
connected to Lord Howe Island (Standard, 1963). 


Basic climatological data for Lord Howe Island is presented in Fig. 2. 


THE PRESENT STATUS OF THE RECENT REPTILES RECORDED FROM 
Lorp Howe ISLAND 


The only recent terrestrial reptiles recorded from Lord Howe Island are 
three species of lizards, only one of which—the skink Leiolopisma lichenigera 
(O’Shaughnessy )—is believed tobeendemic. The other two species are the geckos 
Phyllodactylus guenthert Boulenger and Gehyra oceanica (Lesson). The former 
has been recorded only from Lord Howe Island, Norfolk Island and north- 
western Australia, whereas the latter is widely distributed throughout the 
south-west Pacific region. 

Loveridge (1934) considered that Leiolopisma lichenigera (O’Shaughnessy), 
described originally from Lord Howe Island, was conspecific with Leiolopisma 
cuprea (Gray). The latter species was described from a specimen without 
locality data. The validity of Loveridge’s action is discussed below. 


The first record of the gecko Gehyra oceanica from Lord Howe Island is that 
of Boulenger (1885), who listed a male specimen in the Vritisu Museum (Natural 
History) from ‘“‘ Lord Howe’s Island ’’, collected by J. Macgillivray. Subse- 
quently Etheridge (1889) and Ogilby (1889) included Gehyra oceanica in the 
fauna of Lord Howe Island, but apparently based their record solely on the 
Macgillivray specimen listed by Boulenger. The Australian Museum expedition 
about which Etheridge and Ogilby were writing did not obtain specimens of 
this species. 

Paramonov (1960), presumably following Etheridge and Ogilby, included 
Gehyra oceanica in the Lord Howe Island fauna, while Mitchell (1965) records 
this species from Lord Howe Island on the basis of Zietz’s (1920) record and 
‘“ specimens in the Australian Museum collection”. Zietz appears to have 
based his record on the British Museum specimen recorded by Boulenger, while 


PROCEEDINGS OF THE LINNEAN SocretTy or New SoutH WaAtxgsS, VoL. 96, Part 1 


H. G. COGGER Dik 


there is one specimen (no. 4931) in the Australian Museum which is ostensibly 
a Gehyra oceanica from Lord Howe Island. However, as this specimen has 
apparently been lost (it was collected in the late 1800’s) and as its identification 
and locality are based on a series of ambiguous ditto marks below entries for 
Gehyra oceanica from Fiji and Phyllodactylus guentheri from Lord Howe Island, 
I regard the record as suspect. 


As Gehyra oceanica has been recorded from New Zealand (McCann, 1955) 
it might seem that excessive pains are being taken to discredit the Lord Howe 
Island record for this species. However, as it is normally common wherever 
it occurs, its apparent absence from Lord Howe Island appeared to the author 
to be sufficient grounds for questioning the validity of the earlier records. 


Maegillivray, as naturalist on the voyage of the Herald, spent some months 
on Lord Howe Island (Macgillivray, 1852-55). However, as there were, at the 
time of the Heraild’s voyage, three other islands known as ‘‘ Lord Howe Island ”’ 
in the area covered by the ship’s travels (one, now known as Ontong Java in the 
Solomon Islands; one, now known as Mopeha Island, in the Society Islands ; 
and one in the Santa Cruz Islands), a request was made to Miss A. G. C. Grandison 
of the Herpetology Department of the British Museum (Natural History) to 
examine the original record of Macgillivray’s specimen. Miss Grandison (in litt.) 
reports: ‘‘ In our collection are four specimens given as coming from (a) Fiji, 
(b) Lord Howe Island, (c) two specimens from Fiji; but reference to the original 
register indicates that the specimen (a) has no locality, was part of the Macgil- 
livray collection but was purchased from Cuming. The specimen (b) is given as 
having been collected on the Isle of Pines (New Caledonia) and not Lord Howe 
Island and only the two specimens (c) seem to be correct in their locality.” 
Hence the Macgillivray specimen, on which is based virtually all Lord Howe 
Island records of this species, did not in fact come from that island. For this 
reason it seems advisable to remove Gehyra oceanica from the list of the island’s 
fauna until such time as it is clearly shown to occur there. The remaining two 
Species are redescribed below; S.D. represents the standard deviation of the 
sample. 


Leiolopisma lichenigera (O’Shaughnessy) 
(Plate Om, c) 


Mocoa lichenigera O’Shaughnessy, 1874. Ann. Mag. Nat. Hist., Series 4,13: 298. 


he : 55.8.16.12 
Holotype—British Museum (Natural History) No. 1946.8.16.27 from Lord Howe 


Island, N.S.W. 


Description: Habitus moderate; mean adult snout-vent length 78 mm., 
maximum recorded snout-vent length 96 mm. (2, R26037) ; the snout-vent/axilla 
groin ratio averages 1-93 (range 1 - 74-2 -33, S.D.=0-079) ; limbs well-developed, 
the snout-vent/forelimb length ratio averages 3-90 (range 3 -50—4 -35,S.D.=0-192) 
while the snout-vent/hindlimb length ratio averages 2-99 (range 2-51-3-48, 
S.D.=0-167); the adpressed limbs fail to meet; snout moderately angular ; 
the head is distinctly swollen posteriorly due to the well-developed muscles in 
the quadrate region ; lower eyelid with an undivided transparent disc ; supra- 
nasals absent; nostril pierced in a partly divided nasal, a deep groove dividing 
the scale posterior to the nostril ; frontonasal much broader than long, in contact 
with the rostral anteriorly and the frontal posteriorly ; prefrontals widely to 
narrowly separated ; frontal only slightly longer than broad, about as long as 
frontoparietal and interparietal together, and in contact with the two anterior 
supraoculars ; 4 supraoculars, although in one specimen the two anterior supra- 
oculars on one side are partially fused ; 2 loreals; 7 supralabials, the 4th, 5th 
and 6th below the eye but totally or partially excluded from the granules of the 
lower eyelid by a series of small suboculars ; 5-6 infralabials ; either a single 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Watss, Vou. 96, Part 1 


28 REPTILES OF LORD HOWE ISLAND 


pair of nuchals or no discernible nuchals (of 44 specimens examined for this 
character, 59°% possessed one pair of enlarged nuchal shields, 99% possessed 
an enlarged nuchal on one side only, and 32° lacked any enlarged nuchals) ; 
36-46 scale rows around the middle of the body (mean 42-7, S.D.=1-96) ; 
86-109 rows of scales along the mid-dorsal line from the parietals to the base 
of the tail (mean 99-4, S.D.=3-90) ; 15-21 lamellae under the fourth toe (mean 
18-0, S.D.=1-05) ; ear-opening vertically elliptic, its long axis from 14—2 times 
the diameter of the transparent disc in the lower eyelid. 


Dorsal surface, including the head and limbs, rich bronzy brown to olive 
with scattered dark brown and light brown flecks and streaks. The darker 
flecks tend to disappear in adult animals, so that the lighter ones predominate 
in large individuals. A narrow (usually less than 1 scale in width) brown band 
on either side commencing at the nostril and extending below the canthus to the 
eye. This band continues behind the eye and extends above the ear and along 
the dorso-lateral surface of the body to the tail, and, where the latter is original, 
along the length of the tail. 


This dark dorso-lateral band may continue with equal intensity to the base 
of the tail, but in some specimens it becomes obscure posterior to a point about 
midway between snout and vent. 

The narrow dark dorso-lateral band is bordered below by a dark brown area, 
about 3—4 scales in width, in which scattered groups of light-coloured scales form 
irregular blotches or indistinct oblique bars that are directed downwards and 
backwards. This region fades into the light grey or brown ventro-lateral colour, 
which in turn merges with the cream ventral surface. The latter, in living 
specimens, is often brilliant orange or lemon-yellow. The throat, especially in 
the labial region, is usually flecked with light grey or brown. The upper labials, 
especially in juveniles, may be alternately barred with brown and light grey. 
In life, regenerated tails are usually bright orange. 


Distribution: Leiolopisma lichenigera is known only from Lord Howe Island 
and the small islands lying off-shore from it (including Ball’s Pyramid). 


Material Examined: Australian Museum, Sydney: R26033-R26037, R26057, 
R26074, R26086, Roach Islet, Lord Howe Island; R18919-R18920, 
R18979-R18983, R26048-R26052, Rabbit Island, Lord Howe Island; R2546, 
R7923-R7928, R10942-R10946, R26058-R26071, Lord Howe Island ; 
R29052—R29057, Ball’s Pyramid. Dominion Museum, Wellington: R599, 
Lord Howe Island ; R768, north end of Lord Howe Island. Museum of Com- 
parative Zoology, Harvard: 35459-62, 35464-69, 93776-80, Lord Howe Island. 


Remarks : Loveridge (1934) placed L. lichenigera in the synonymy of Leiolopisma 
cuprea (Gray), a species whose type locality is unknown. Boulenger (1887), 
who had the holotypes of both species before him, differentiated the two species 
on the following characters : 


lichenigera cuprea 
4 supraoculars 3 supraoculars 
42 mid-body scale rows 36 mid-body scale rows 


Loveridge had a series of 18 specimens, all of which possessed 4 supraoculars 
and with mid-body scale row counts ranging from 36-46 (mean 42). He 
suggested that the 3 supraoculars in the type of cuwprea were abnormal. As 
L. lichinigera is known only from Lord Howe Island, Loveridge’s action implied 
that the type of L. cuprea came from this locality. 


Of the 69 specimens listed above, only one shows any deviation from the 
four supraoculars count. This specimen (R18919) has only 3 supraoculars 
(caused by partial fusion of the first and second) on the right-hand side. The 
mid-body scale row count varies from 36 to 46, averaging 42-7. 


PROCEEDINGS OF THE LINNEAN SocreTy or New SoutH WaAtEs, Vou. 96, Part 1 


H. G. COGGER 29 


In the scineid genus Hmoia, Brown (1953, 1954) made good use of the number 
of scale rows along the mid-dorsal line, from parietals to tail base, as a taxonomic 
character. This count was made on all of the above specimens of L. lichenigera 
and averaged 99-7 (range 85-109, S.D. of sample 3-93). 


The holotype of L. cuprea has three supraoculars on each side, 36 rows of 
scales around the middle of the body and about 84 rows of scales from the 
parietals to the base of the tail (damaged parietal region prevents a precise 
count). It is in poor condition, being so badly discoloured that the original 
colour and pattern are not discernible. The nostril is pierced in an otherwise 
immaculate nasal, whereas the nasal is strongly grooved or divided posteriorly 
in all specimens of ZL. lichenigera examined. 


A specimen (M.C.Z. 35460) cited by Loveridge (1934) is the only one found 
to possess a mid-body scale row count below 40. This specimen has only 36 rows 
of scales around the middle of the body, and has only 86 rows of scales from 
parietals to base of tail. Loveridge considered that this exceptional individual 
bridged the gap between cuprea and lichenigera, but as the latter is known only 
from the one island locality it seems unlikely that either geographic or individual 
variation could account for the differences between this species and the type 
specimen of cuprea. 


In view of this decision to regard LZ. cuprea and L. lichenigera as distinct 
Species, an attempt was made to determine with what other known species 
L. cwprea might be conspecific. In its colour, pattern, the undivided nasal and 
the shape and disposition of many head scales (notably the labials, loreals and 
preoculars) it appears to be most closely allied to the Leiolopismas of New 
Zealand. However, it differs from the described species in enough features 
(especially its supraocular count) to cast considerable doubt as to its relationships. 
It is therefore relegated to the status of a nomen dubium. 


Phyllodactylus guenthert (Boulenger) 
(Plate I, D) 
Phyllodactylus guenthert Boulenger, 1885. Cat. Iiz. Brit. Mus., 1: 90, pl. 7, 
fig. 3. 
Syntypes: (5), British Museum (Natural History) from Champion Bay, N.W. 
Australia (67.2.19.18), Lord Howe’s Island (56.7.7.19) and Norfolk Island 
(62.1.6.29-31). 


Description: Habitus stout ; mean adult snout-vent length 71-2 mm., maximum 
recorded snout-vent length 91 mm. (R29193); limbs well developed; the 
snout-vent/forelimb length ratio averages 3:09 (range 2-66-3-70, S.D.=0-223) 
while the snout-vent/hindlimb length ratio averages 2-47 (range 2-00-2-75, 
S.D.=0-155); the adpressed limbs overlap; head relatively deep; canthus 
rounded ; eye moderate, its diameter equal to about half the distance from eye 
to tip of snout; rostral rectangular, about twice as wide as deep and lacking a 
median crease ; nostril directed backwards and upwards and surrounded by the 
rostral, first labial and four other scales, the upper one or two of which are 
considerably enlarged ; several small scales between these enlarged ‘‘ supra- 
nasals ’; scales on top and sides of snout larger than dorsals; but the latter 
and the scales on the head are subequal; 7-9 supralabials; 7-9 infralabials ; 
the anterior labials are largest, and all are much larger than the adjacent scales ; 
mental roughly pentagonal, about the same size as adjacent infralabials ; anterior 
and lateral gular scales much larger than the remainder ; a distinct gular fold : 
ear-opening relatively small, its maximum diameter about two-thirds that of the 
eye; dorsal and lateral surfaces of body covered with small, juxtaposed conical 
scales which are much smaller than the flat, slightly-imbricate ventrals ; more 
than 10 scales in the middle of the back in a distance equal to the diameter of 
the eye; number of scale rows around middle of body averages 138-4 (range 


PROCEEDINGS OF THE LINNEAN Society oF New Souru Wates, Vou. 96, Part 1 


30 REPTILES OF LORD HOWE ISLAND 


119-160, S.D.=7-87) ; limbs covered above with conical scales somewhat larger 
than those on the dorsum ; lower surfaces of hmbs with flat, slightly imbricate 
scales like those on the venter; digits moderately dilated, the distal expansion 
of the 4th toe greatest, its width about equal to the maximum diameter of the 
ear-opening ; each digit with a pair of enlarged pads or lamellae distally, between 
which is a large claw; the sheath from which each claw projects extends to 
the edge of the distal lamellae or beyond; there are 9-12 undivided lamellae 
under the 4th toe (mean 10-23, S.D.=0-815); these are largest basally and 
are separated from the enlarged distal lamellae by from 3-7 rows of small scales ; 
tail covered with flat, slightly imbricate scales which are arranged in fairly 
regular rings and are largest ventrally ; the latter are about twice as large as 
those on the upper surface of the tail, these latter more or less equaling in size 
the scales on the venter; preanal and femoral pores absent; males with a 
swollen post-cloacal region on each side of which is a cluster of 3-4 shghtly 
enlarged rounded tubercles. 


Colour and pattern extremely variable. Dorsal surface medium grey or 
brown (the latter colour predominates in preserved specimens) usually with a 
dull orange (in life) or light brown vertebral stripe which extends from the nape 
to the base of the tail. This stripe usually divides a series of irregular, transverse 
dark grey or dark brown bands which typically number six between the nape 
and the base of the tail. In many specimens this vertebral stripe may be 
indistinct or absent, while the irregular transverse bands may break up to such 
an extent that the dorsal surface is simply variegated with light and dark grey 
or brown. The head is often darker than the ground colour of the body. Labials 
mottled with light grey or brown ; sides and upper surfaces of the limbs are usually 
grey or brown mottled with irregular darker markings, while the ventral surface 
is grey or off-white, each individual scale peppered with dark grey. The scales 
of the throat are more richly peppered than those of the body. 


The tail, where original, is usually alternately banded with irregular dark and 
light grey or brown, the bands not extending on to the ventral surface ; in some 
specimens these bands break up to form irregular mottling. Regenerated 
tails are usually light brown or grey with scattered darker scales. Underside 
of tail light grey. 

No major differences in colour and pattern can be observed between the 
Lord Howe Island and Norfolk Island populations. However, there is a definite 
tendency for the darker transverse markings to be narrower and less conspicuous 
in the Norfolk Island specimens, while old adults from the latter locality tend 
to have the pattern reduced to scattered dark flecks or spots. 


The colour and pattern of hatchlings from Lord Howe Island appear to be 
fairly constant. The dorsal ground colour is rich olive-brown, speckled with 
darker brown; there is a light salmon-coloured patch over each eye and an 
indistinct, broken vertebral stripe of a rich salmon colour; the tail, especially 
distally, is rich salmon with dark brown bands ; the venter is white with numerous 
grey flecks ; the ventral surface of the tail is salmon-coloured. 


Distribution: Phyllodactylus guenthert is known from Norfolk Island and Lord 
Howe Island and their small off-shore islands. In his original description 
Boulenger lists one of the type series from ‘‘ Champion Bay, N.W. coast of 
Australia’, but this record appears to be in error. Champion Bay lies within 
the known range of Phyllodactylus marmoratus and is far removed from the other 
localities in which P. guentheri is known to occur. No specimens of the latter 
have subsequently been recorded from mainland Australia. However, Miss 
Grandison has checked the original data associated with this specimen and can 
find no reason for doubting its validity. 

Material Examined: Australian Museum, Sydney: 4930, R360, R3350, R4542, 
R5581, R5583-4, R8043, R26072-3, Lord Howe Island ; R26038-47, R26085, 


PROCEEDINGS OF THE LINNEAN Soctety or NEw SoutH WALES, Vou. 96, Part 1 


H. G. COGGER aul 


Roach Island, Lord Howe Island; R18984-6, R19298-9, R26053-6, Rabbit 
Island, Lord Howe Island; R4270-3, R4275-7, R4919, R5594, R19301, 
R29192-4, Norfolk Island; R29058-9, Ball’s Pyramid, Lord Howe Island. 
Dominion Museum, Wellington: R1-8, R780-2, Norfolk Island. 


Remarks: This species has long been in the synonymy of Phyllodactylus 
marmoratus, the only other Australian member of this genus (Loveridge, 1934). 
However, P. guenthert is very distinct from P. marmoratus, and can be readily 
distinguished from the latter : 

Phyllodactylus marmoratus : Width of distal expansion of 4th toe at least twice 
the maximum diameter of the ear-openings ; scales relatively large and regular ; 
84-96 mid-body scale rows (mean of 21 specimens 90-4, S.D.=3-7); sheath 
of claw on each digit does not extend to edge of distal lamellae (Fig. 3). 


Fig. 3. Lateral and dorsal views of distal portions of the fourth toes of Phyllodactylus 
guenthert (upper) and P. marmoratus (lower). 


Phyllodactylus guenthert: Width of distal expansion of 4th toe about equal to 
the maximum diameter of the ear-opening ; scales on most parts of body small, 
tubercular ; 119-160 mid-body scale rows (mean 138-4); sheath of claw of each 
foot extends to edge of distal lamellae or beyond (Fig. 3). 


Unfortunately the material available for study is inadequate for an effective 
analysis of intraspecific variation in meristic and other characters. Nevertheless, 
there are indications that the populations on the different islands do differ from 
one another at a measurable level. For example, the two specimens from Ball’s 
Pyramid have much longer limbs than specimens from other localities. The 
data in Table 1 are presented, therefore, with a view to indicating the potential 


PROCEEDINGS OF THE LINNEAN Socrpty or New SoutH Watzs, Vou. 96, Part 1 


32 REPTILES OF LORD HOWE ISLAND 


value of using these island populations for micro-evolutionary studies. There 
seems little doubt that barriers to dispersal and interchange between the various 
island populations are so effective that gene flow is reduced to a mere trickle, 
if any, and that each island population is evolving almost entirely independently 
of the others. 


TABLE | 
Mid-body Lamellae Snout-vent/ Snout-vent/ Snout-vent/ 
Scale under Axilla- Forelimb Hindlimb 
Rows Fourth Toe groin 
Ball’s Pyramid 145-147 9-11 2:29-2:47 2-58-2-71 2-00-2-28 
(2) (146-0) (10-0) (2-38) (2:64) (2-14) 
Roach Island 135-160 9-10 2-16—-2-44 2:75-3:04 2:27-2:54 
(10) (146-3) (9-9) (2-32) (2-89) (2-39) 
§8.D.=7:44 §$.D.=0-38 8.D.=0:064 S8.D.=0-102 S.D.=0-058 
Rabbit Island 136-153 10-12 2:17-2:43 2-91-3-70 2-41-2-74 
(9) (142-2) (11-0) (2:27) (3:27) (2-58) 
§8.D.=6-97 S.D.=0:059 S.D.=0-261 S.D.=0-233 
Lord Howe Island”) 137-147 10 2-18—-2-90 2-66—3-17 2-28-2-58 
(8) (139-2) (10-0) (2-40) (2-94) (2-44) 
8.D.=4:-38 8.D.=0-225 S.D.=0-154 §.D.=0-088 
Lord Howe Island‘) 135-160 9-12 2-16-2-90 2:58-3:70 2-00-2-74 
(28) (143-7) (10-14) (2-33) (3-01) (2-45) 
§8.D.=6:79 S.D.=0-77 S.D.=0-:132 S.D.=0:255 §.D.=0-294 
Norfolk Island 119-140 9-12 2:00-2:51 2-84-3-59 2-14-2-75 
(24) (133-7) (10-24) (2-19) (3-13) (2-50) 
§8.D.=4:82 S8.D.=0:90 S.D.=0-111 S8.D.=0-181 S.D.=0-163 


@) Main island only. 
) Lord Howe Island and off-shore island specimens pooled. 


The relationship between P. guentheri and P. affinis (the latter known only 
from the New Hebrides) is obscure. Miss Grandison kindly examined the 
holotype of P. affinits and found numerous differences in scalation between the 
two species. It would therefore seem advisable to leave any assessment of 
these differences until additional specimens of P. affinis are available. 


LOCAL DISTRIBUTION AND ECOLOGY 


The first report on the habits of the lizards of Lord Howe Island was that of 
Etheridge (1889), who stated of Phyllodactylus guentheri and Lygosoma lichenigera 
that they “‘...may be sought for on dry stony ground, under stones, amongst 
dry leaves, and at the feet of low scrubby trees.” Ogilby (1889) states simply 
that both species are “‘. . . much more abundant in the vicinity of the coast than 
on the more elevated districts inland.’? Etheridge (1889) goes on to say that 
‘““...80 far as our researches enabled us to judge, Phyllodactylus guentheri is 
the commonest, the distribution of this and the other species (ZL. lichenigeruwm) 
being very general, not only on the main island, but also on Goat (=Rabbit) 
Island and the Admiralty Islets.” 

No further general account of the reptiles was given until that of Paramonov 
(1960), who, although he did not mention Etheridge’s paper, nevertheless quoted 
the latter precisely. However, in doing so he. made the error of applying 
Etheridge’s comments on habitat to all three species, instead of the two species 
to which Etheridge originally and correctly referred. For reasons outlined 
above the inclusion of Gehyra oceanica was probably in error. 


PROCEEDINGS OF THE LINNEAN Socrety or New SoutH WALES, Vou. 96, Part 1 


H. G. COGGER 33 


It is significant that despite the fact that no zoologist had previously visited 
Lord Howe Island for the express purpose of collecting reptiles, small collections 
have nevertheless been obtained apparently with little difficulty. The Australian 
Museum possesses several small series of specimens collected at various times 
between 1888 and 1963. Except for the series collected in 1962-63 from Rabbit 
Island, all earlier specimens have the locality of ‘‘ Lord Howe Island’. As the 
prime reason for the visit of an Australian Museum party in 1909 was to obtain 
bird specimens from the Admiralty Islets for a Museum exhibit, it seems likely 
that at least some of our lizards were taken from those islets and not from the 
main island. 


Loveridge (1934) states that aseries of specimens of L. cuprea (=L. lichenigera) 
in the Museum of Comparative Zoology was collected by Mr. R. Baxter in 1932. 
These specimens were taken when they fell into a drum set into the ground and 
baited with fish oil, and K. R. Hindwood (in litt.) reports that Baxter informed 
him that these specimens were taken on Rabbit Island after unsuccessful attempts 
to collect lizards on the main island. A specimen (R768) of L. lichenigera in 
the Dominion Museum, Wellington, collected by W. R. B. Oliver in 1913, has 
the notation ‘‘ north end Lord Howe Island ”’. 


From this scant evidence there seems little doubt that both species— 
Phyllodactylus guenthert and Letolopisma lichenigera—were once common and 
easily procured on the main island, as well as on the off-shore islands, and it was 
with this background information that the writer set out to assess the present 
distribution of reptiles on Lord Howe Island. 


The results were disappointing. Despite a search of all major available 
habitats, especially those in which earlier workers had reported finding lizards, 
not a single specimen was obtained. Although some habitats, such as the leafy 
heads of the Howea palms, were virtually inaccessible, all terrestrial habitats 
were examined. Three major saxicoline environments are present on the main 
island : 
(1) Covering the greater portion of the island are outcrops of basaltic 
volcanic rocks, in the form of either solid rock faces or boulder slopes. All of 
the major topographic features are made up of this rock, as are all of the off-shore 
islands. 
(2) Recent sediments, which form a stratified ‘“‘ sandstone’, outcrop in 
several places, the most prominent being above Ned’s Beach on the eastern side 
of the island. 
(3) The only other rock consists of large clumps of old ‘ beach rock ”’, 
which may be loosely or firmly embedded in soil, and which outcrop in several 
places on the lower parts of the island. 
The first of these three rocky environments corresponds with the ‘“‘ North 
Ridge Voleanics ’’ and ‘‘ Mt. Lidgbird Voleanics ”’ of Standard’s (1963) geological 
map of the island. The second corresponds with his ‘‘ Ned’s Beach Calcarenite ” 
and the third with his ‘“‘ Alluvium, including beach and talus deposits’’. The 
age and distribution of each of these are shown on Standard’s map. 
On the main island a search of the basalt rock faces and slopes failed to 
reveal a single lizard. Collecting efforts were concentrated at four points : 
(1) The boulder slopes below Mt. Lidgbird, immediately behind Salmon 
Beach. 

(2) The slopes of Dawson Ridge, between Old Settlement Bay and North 
Bay. 

(3) The lower, eastern slopes of Malabar. 

(4) The northern slopes of Intermediate Hill. 


Also, although the search was far from exhaustive, a search was nonetheless 
carried out along the track from Salmon Beach to the summit of Mount Gower 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WaAtLEs, Vou. Part 96, 1 


34 REPTILES OF LORD HOWE ISLAND 


and in the ‘‘ moss forest ’? which occupies several hundred acres on the summit 
of that mountain. 


All of these searches failed to find any trace of lizards, either among rocks 
or in or under fallen logs. No sloughs were found ; nor eggs or egg shells. 


Following reported sightings of lizards, the area behind Ned’s Beach was 
also searched carefully, especially in the stratified ‘‘ sandstone ’’ which makes up 
much of the headland (Stevens Point) between Ned’s Beach and Middle Beach. 
No lizards were found. 


During the time that this work was being carried out, a cyclone prevented 
the author from reaching any of the off-shore islands. As soon as the weather 
cleared a visit was made to Rabbit Island, where both Phyllodactylus guentheri 
and Leiolopisma lichenigera were found commonly under loose basalt boulders. 
Small slopes of these loose boulders occurred mainly on the leeward (northern) 
side of the island, especially around the bases of low, windswept trees. On the 
windward side of the island few loose boulders occurred but an examination of 
large crevices in outcrops of rock revealed traces (a mummified body and sloughed 
skins) of both species. 


As the island is so small, and as the Australian Museum already possessed 
samples from Rabbit Island, only a small number of specimens were actually 
taken. Although unlikely, it is nevertheless quite possible that the removal of 
large numbers of lizards could have a decimating effect on the total population. 
Of the lizards found, the ratio of L. lichenigera to P. guentheri was 4:1. 


Subsequently, attempts were made to land on two of the other off-shore 
islands—Mutton Bird Island, off the eastern shore, and Roach Island, the largest 
of the Admiralty group of islands off the southern point of the main island. Both 
of these islands are surrounded by rocky shores, and although heavy seas 
prevented a landing on Mutton Bird Island, the author was put ashore on 
Admiralty Islet, where collecting was carried out for about an hour before rising 
seas forced a hurried and wet return to the boat. During this hour 19 specimens 
were collected among large basalt boulders, in the ratio of 11 P. guentheri to 
8 L. lichenigera. Both species were evidently extremely abundant. 


It should be stressed that the lizards taken on Rabbit Island and on Admiralty 
Islet were found in a habitat which had proved completely barren of lizards on 
the main island. 


About half-way through the author’s visit to Lord Howe Island he was 
presented with a live specimen of L. lichenigera by Mr. R. Payton, whose cat 
had apparently caught the lizard in the vicinity of the Payton house. The 
latter is situated behind Lagoon Beach, just below Signal Point. In the following 
week the cat produced three more specimens of the same species. The area in 
which these lizards were taken consists of dense Howea palm forest with leaf- 
litter and scattered clumps of weathered ‘“‘ sandstone ’’ embedded in the soil. 
The nature and origin of this rock is discussed below. 


In a similar area, at the base of Malabar, the author searched among similar 
‘‘ sandstone ”’ boulders and found a specimen of P. guentheri which entered one 
of the numerous finger-sized tunnels in a massive clump of the rock. The gecko 
could not be extricated from the rock but in some of the shallow holes clusters 
of eggs were found, some of which contained advanced living embryos of P. 
guenthert. 


Subsequently, a sledge-hammer and crowbar were used unsuccessfully in 
an attempt to break up some of these sandstone clumps in the areas below Malabar 
and behind Lagoon Beach. Only the thinner surface layers of the stone could be 
broken up, leaving intact the massive bases, with their network of tunnels. 
No adult lizards were obtained but some hundreds of old and new gecko eggs 
were found, some with well-developed living embryos. 


PROCEEDINGS OF THE LINNEAN Society or New SourH WALES, Vou. 96, Part 1 


H. G. COGGER 35 


This habitat warrants description in greater detail, as it is apparently 
the only habitat on the main island in which both species of lizards now occur. 
It consists of large irregularly-shaped boulders of honeycombed ‘‘ rock”? which 
may be either superficially or deeply embedded in the soil (Pl. 1, D). 


These boulders almost invariably outcrop in dense Howea palm forest, in 
which there is a thick mat of humus formed by palm and other leaf-litter. The 
matrix of the boulders consists of a “‘ sandstone’ or conglomerate sediment, 
and there seems little doubt that the outcrops are of weathered ‘‘ beach rock ”’, 
a common formation of many coral reefs and the structure and origin of which 
is discussed fully by Wiens (1962). According to Standard (1963) these outcrops 
represent recent beach alluvium. 

Of special significance in the ecology of the lizards on Lord Howe Island is 
the mode of weathering of these alluvial boulders. Their surfaces are deeply 
pitted with small holes, averaging only 1-2 cm. in diameter. These are the 
entrances to tunnels which enter deeply into the matrix of the rock. Such 
tunnels may coalesce to form larger chambers, as shown in Pl. U, A, B. The 
origin of these tunnels is uncertain, but they so closely resemble in structure the 
‘“‘ subaerial solution potholes ”’ observed in some limestones (Cotton, 1948, p. 448) 
that their origin might well be identical, especially as buried portions of these 
boulders are as strongly weathered as the exposed portions. Lord Howe Island 
has a relatively high average annual rainfall (68 inches) and the deeply-shaded 
floor of the Howea forests is normally quite moist. The exposed surfaces of the 
outcropping beach rock are covered with a layer of algae which imparts a 
distinctive green colour to the rock. 


The role of the weathered tunnels or potholes in the ecology of the two lizards 
on Lord Howe Island seems to be that they provide a niche which is safe from 
virtually all predators, for the size of the openings precludes the entry of any 
of the island’s predatory vertebrates. The tunnels offer an ideal site for the 
safe deposition of eggs (PI. m1, A) while insect food, both in the tunnels and in the 
surrounding leaf-litter, is abundant. The eggs of Phyllodactylus guentheri 
may be found in large numbers within the tunnels, recent clutches being deposited 
upon the hatched, empty shells of earlier seasons (Pl. WH, B). 


Apart from those eggs broken or immediately preserved, a series of five 
intact and apparently fertile eggs were brought back to the Australian Museum. 
There they were placed on paper in a dry glass jar, and left on a laboratory 
bench, where they were subject to a mean temperature of approximately 70° F. 
(range 66° F.-84° F.). As none had hatched by the end of December 1966 
(the eges having been collected in June 1966), they were no longer examined 
regularly and were put to one side. When next examined on 18th January 
1967 two of the eggs had hatched. Both hatchlings were dead, one freshly so 
but it was possible to place the date of hatching within the two-week period 
preceding the 18th. Another egg hatched on 8th February, another on 13th 
February and another on 14th March. The mean snout-vent length of hatchlings 
was 30-25 mm. 

The eggs of P. guentheri have thick, brittle calcareous shells which are 
apparently quite soft when laid, for they take the shape of the surfaces with 
which they come into contact when laid. Eggs are often found firmly adhering 
to one another or to the surface of a rock. 

The significance of this long incubation time is not known. As it was not 
possible to know how long the eggs had been laid when collected, the minimum 
incubation time for those eggs collected varied from 30 to 39 weeks. Although 
overwintering of eggs is a relatively common phenomenon in lizards, the observa- 
tions made above are not explicable in normal overwintering terms. The eggs 
were collected in mid-winter (when no gravid females were collected) and 
continued to develop throughout the spring and summer. Indeed, the last egg 


PROCEEDINGS OF THE LINNEAN SocreTy OF NEw SoutH Watss, Von. 96, Part ] 


36 REPTILES OF LORD HOWE ISLAND 


to hatch did so at the end of summer. The temperature of development in the 
laboratory was probably not very different from that experienced by eggs in the 
field during this period. 

From these meagre observations it would be unwise to draw any general 
conclusions regarding incubation times or their roles in the biology and ecology 
of this lizard. These observations do indicate, however, that the reproductive 
habits of Lord Howe Island reptiles may differ radically from those of related 
forms on the Australian mainland. 


On returning to Sydney, the author contacted Mr. David Rootes, a member 
of several climbing parties which attempted to scale Ball’s Pyramid. Mr. 
Rootes advised that one species of lizard had been seen, and produced several 
photographs showing clearly an individual of L. lichenigera. This individual 
was observed to roll a tern egg down a rock slope until it broke, then fed on the 
contents. 

Mr. Rootes’ observations were made in November, whereas no lizards were 
active in the open during the author’s visit to Lord Howe Island in winter (June). 


SUMMARY 


(1) The recent reptile fauna of Lord Howe Island comprises two species of 
lizards, the gecko Phyllodactylus guenthert and the endemic skink Leiolopisma 
lichenigera. The latter is resurrected from the synonymy of Leiolopisma cuprea, 
which is in turn relegated to the status of a nomen dubium. It has been shown 
that earlier records of the gecko Gehyra oceanica from Lord Howe Island were 
probably in error. 

(2) On the small islands lying off-shore from Lord Howe Island the lizards 
occur commonly in the rocky habitat described by earlier workers. On the main 
island, however, both species appear to have virtually disappeared from the 
habitat in which they were once common and are now to be found only in a very 
restricted and specialized habitat consisting of weathered boulders of ‘ sand- 
stone ”’ or “‘ beach rock’. It seems likely that this habitat is the only one which 
offers protection from such introduced predators as rats. 


(3) Slight differences in the proportions of the two lizards on different islands, 
differences in size, and the prolonged incubation period of the eggs of Phyllo- 
dactylus guentheri indicate the potential value of the reptile fauna of Lord Howe 
Island for future ecological and evolutionary studies. 


ADDENDUM 


Since this paper was submitted for publication the types of Phyllodactylus 
guenthert and P. affinis have been examined in the British Museum (Natural 
History). The P. guentheri syntype from Champion Bay is indeed conspecific 
with the other syntypes, and I therefore regard its locality data as incorrect. 
The holotype of P. affinis (B.M.N.H. 55.8.16.5.) appears to be a typical example 
of the Australian Phyllodactylus marmoratus, for which reason I suggest that its 
type locality (New Hebrides) is also incorrect. 

Further to my comments on the ecology of Leiolopisma lichenigera, Dr. 
H. Recher reports (pers. comm.) that in summer this species is abundant and 
active at night among supralittoral rocks and other ground cover on Rabbit 
Island. 


ACKNOWLEDGEMENTS 
The author is grateful to the Trustees of the Australian Museum and the 
Museum’s Director, Dr. F. H. Talbot, for making possible a visit to Lord Howe 
Island, and to the Lord Howe Island Board for permission to undertake the 
work here reported upon. Mrs. Kirby (then of ‘‘ Pine Trees’), Mr. R. Payton, 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Watss, Vou. 96, Part 1 


4 


H. G. COGGER 37 


Mr. EK. Rhodes, Mr. KR. Schick and Mr. C. Wilson were especially kind and helpful 
during the author’s stay on the island. Miss A. G. C. Grandison of the British 
Museum (Natural History), Dr. Ernest Williams of the Museum of Comparative 
Zoology at Harvard University, and Dr. J. Moreland of the Dominion Museum, 
Wellington, kindly examined and/or lent specimens under their charge. 


References 


Brown, W. C., 1953.—Results of the Archbold Expeditions. No. 69. A Review of New Guinea 
Lizards Allied to Hmoia baudini and E'moia physicae. Amer. Mus. Novit.. No. 1627, pp. 1-25. 
, 1954.—Notes on several lizards of the genus Hmoia, with descriptions of new species 
from the Solomon Islands. f&ieldiana: Zoology, 34 (25): 263-276. 
BouLencer, G. A., 1885.—Cat. Liz. Brit. Mus. (N.H.), 1, British Museum (N.H.), London. 
, 1887.—Cat. Liz. Brit. Mus. (N.H.), 3, British Museum (N.H.), London. 

Cotton, C. A., 1948.—Landscape. As developed by the processes of normal erosion. 2nd EKd., 
Whitcombe and Tombs Ltd., Christchurch, pp. 1-509. 

ETHERIDGE, R., 1889.—The General Zoology of Lord Howe Island, in Lord Howe Island. Its 
Zoology, Geology and Physical Characters. Memoirs Aust. Mus., 2 (1): 3-42. 

GurRNEY, A. B., 1947.—Notes on some remarkable Australian walkingsticks, including a synopsis 
of the genus Hatatosoma (Orthoptera: Phasmatidae). Ann. ent. Soc. Amer., 40: 373-396. 

Hinpwoop, K. A., 1940.—The birds of Lord Howe Island. Hmu, 40: 1-86. 

, and CUNNINGHAM, J. M., 1950.—Notes on the birds of Lord Howe Island. Hmu, 
50: 23-35. 

Loverinc#, A., 1934.—Australian Reptiles in the Museum of Comparative Zoology, Cambridge, 
Massachusetts. Bull. Mus. Comp. Zool., Harv., 77 (6): 243-383. 

MacGintivray, J., 1852—55.—Voyage of H.M.S. Herald, under command of Captain H. Mangles 
Denham R.N. being Private Journal kept by John MacGillivray, Naturalist. Mss. No. 23, 
Admiralty Library, London. 

McAtrint, D. K., 1966.—Rediscovery of Lord Howe Island Phasmid. Aust. ent. Soc. News 
Bull., 2 (3): 71. 

McCann, C., 1955.—The Lizards of New Zealand. Gekkonidae and Scincidae. Bull. Dom. 
Mus., 17: i-vin, 1-127. : 

McCuttoca, A. R., 1921.—Lord Howe Island—A Naturalist’s Paradise. Aust. Mus. Mazg., 
1 (2): 31-47. 

McKean, J. L., and Hinpwoop, K. A., 1965.—Additional Notes on the Birds of Lord Howe 
Island. Hmu, 64 (2): 79-97. 

MircHetyt, F. J., 1965.—Australian geckos assigned to the genus Gehyra Gray (Reptilia : 
Gekkonidae). Senck. biol., 46 (4): 287-319. 

Nicuouis, Max, 1938.—Lord Howe Island, 1788-1938. George M. Dash, Sydney, pp. 1-76. 

OcitBy, J. D., 1889.—The Reptiles and Fishes of Lord Howe Island, 7m Lord Howe Island. Its 
Zoology. Geology and Physical Characters. Memoirs Aust. Mus., 2 (3): 51-74. 

OuiveR, W. R. B., 1917.—The Vegetation and Flora of Lord Howe Island. Trans. N. Zeal. 
Instit., 49: 94-161. 

Paramonov, S. J., 1958.—Lord Howe Island, a Riddle of the Pacific. Pacific Science, 12 (1): 
82-91. 

, 1960.—Lord Howe Island, a Riddle of the Pacific. Part II. Pacific Science, 14 (1): 
75-85. 
, 1963.—Lord Howe Island, a Riddle of the Pacific. Pacific Science, 17 (3): 361-373. 

Putte, ARTHUR, 1789.—The Voyage of Governor Phillip to Botany Bay. ..compiled from 
Authentic Papers...etc. John Stockdale, London. 

Pops, HE. C., 1949.—A Naturalist on Lord Howe Island, Part I. Wald Infe, 12 (11): 549-554. 

, 1959.—Lord Howe Island, Ch. 10 in The Great Barrier Reef and Adjacent Isles (by 
K. Gillett and F. McNeill), Coral Press, Sydney, pp. 141-159. 

, 1960.—The Natural History of Lord Howe Island. Aust. Mus. Mag., 13 (7): 
207-210. 

Squirss, D. F., 1963.—Carbon-14 Dating of the Fossil Dune Sequence, Lord Howe Island. Aust. 
J. Science, 25 (9): 412-413. 

STANDARD, J. C., 1961.—Submarine Geology of the Tasman Sea. Bull. Geol. Soc. Amer., 72: 
1777-1788. 

, 1963.—Geology of Lord Howe Island. Jour. and Proc. Roy. Soc. N.S.W., 96: 
107-121. 

Wiens, H. J., 1962.—Atoll Environment and Ecology. Yale University Press, New Haven, 
pp. 1-xxil, 1-532. 

Zierz, F. R., 1920.—Catalogue of Australian Lizards. Rec. S. Austral. Mus., 1: 181-228. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SourH WateEs, Vou. 96, Part 1 


38 REPTILES OF LORD HOWE ISLAND 


EXPLANATION OF PLATES 


PLATE I 
A. Lord Howe Island viewed from Malabar, showing Mt. Lidgbird (left) and Mt. Gower. 
B. Northern, low-lying parts of Lord Howe Island viewed from the summit of Mt. Gower ; Rabbit 
Island is seen lying in the lagoon while the Admiralty Islands are in the right upper corner. 
C. Rabbit Island from Lagoon Beach. 
D. Clump of “ beach rock” riddled with tunnels in which the eggs of Phyllodactylus guentheri 
are laid. 
PLATE II 
A and B. Sections of “ beach rock’’ showing successive clutches of eggs of Phyllodactylus 
guenthert laid within tunnels. 
. Lertolopisma lichenigera. 
. Phyllodactylus guentheri. 


va 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WaAtEs, Vou. 96, Part 1 


PLATE I 


Proc. Linn. Soc. N.S.W., Vol. 96, Part 1 


‘A: 

fl | 
‘ . 
E yt 


noe 
YY 
ii 


Proc: Linn. Soc. N.S.W., Vol. 96, Part 1 PLATE II 


AUSTRALIAN SEA STARS OF THE GENUS PATIRIELLA 
(ASTHROIDEA, ASTERINIDAE) 


A. J. DARTNALL 
Tasmanian Museum, Hobart, Tasmania 


(Plates 111, Iv) 


[Accepted for publication 23rd September 19706] 


Synopsis 
Including the two new species described in this paper ten species of Patiriella are known from 
Australia. The Lamarckian species Patiriella evigua is shown to contain a tropical and temperate 
species. A neotype is erected to validate the identity of P. exigua sensu stricto. A key to the 
Australian species of Patiriella is provided. An account of the biogeography of the-genus in 
relation to known marine zoogeographic boundaries is given. 


INTRODUCTION 


The present paper deals with asterinid sea stars of the genus Patiriella 
from Australia. Patiriella comprises a group of sea stars whose members are a 
conspicuous feature of the intertidal fauna of temperate and cool temperate 
shores in the Southern Hemisphere. Although the primary concern here is the 
fauna of Australia, some reference is also made to the asterinid faunas of South 
Africa, the Indo-west-Pacific area and New Zealand. 


Australia and Tasmania possess eight species of Patiriella and two further 
Species are described in this paper. Patiriella exigua (Lamarck) is common to 
South Africa and Australia and a New Zealand species (Patiriella regularis 
(Verrill)) is known as an introduced member of the Tasmanian marine fauna 
(Dartnall, 1969a). 


Abbreviations used in the text are as follows: A.M., Australian Museum, 
Sydney; B.M., British Museum (Natural History), London; M.C.Z., Museum 
of Comparative Zoology, Harvard; M.N., Museum fiir Naturkunde, Berlin ; 
N.M.V., National Museum of Victoria, Melbourne ; Q.M., Queensland Museum, 
Brisbane ; T.M., Tasmanian Museum, Hobart ; U.S.N.M., United States National 
Museum, Washington, D.C.; W.A.M., Western Australian Museum, Perth. 
The material quoted is housed in the institution indicated by the initial preceding 
the registration numbers given. The number of specimens in each lot is indicated 
in parentheses after the registration number. 

& is the greater radius of the specimen measured along the ambulacral 
groove ; 7 is the lesser, interambulacral radius. Body proportions are expressed 
as a ratio 2:7 and the height of the body is given as vh. 


SYSTEMATIC ACCOUNT 
Genus PATIRIELLA Verrill 1913 

Type Species: Asterina (Asteriscus) regularis Verrill, 1867. 

Asterinid sea stars with a plane actinal surface, short rays and a body 
outline varying from a blunt star to nearly pentagonal. Actinal intermediate 
spines rarely more than two, most often paired or single. Oral plates may carry 
one, two or no suboral spines. The inferomarginal plates and their spines form 
a fringing edge to the body and the superomarginal plates are not distinct. 
Abactinal spines are short, granular and bluntly capitate. The carinal row of 


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40 SEA STARS GENUS PATIRIELLA 


abactinal plates is doubly notched to accommodate two rows of papulae. Papuael 
are confined to the abactinal surface, along the rays, on the centre of the disc 
and the interradial, abactinal areas near the disc centre. The plates of the sides 
of the rays and the outermost interradial areas are apapulate. Pedicellariae 
are absent. 


Species of this genus are found on rocky shores in the intertidal zone of 
southern and eastern Australia. Populations of some species may be found on 
mud, sand or gravel and can be taken below tidemarks to about 15 metres and 
occasionally at greater depths. 

The genus Patiriella is most closely related to the predominantly northern 
hemisphere genus Patiria; to Paranepanthia which contains three nominal 
species in Australia, and to part of the heterogeneous genus Asterina from which 
Patiriella was originally removed. 

In a previous account (Dartnall, 1970b) the genus was subdivided into 
‘“ exigua”’, ‘ regularis””’ and “ gunni”’ groups. ‘There are distinct morphological 
differences between these groups; within them are species of very similar 
morphology whose isolation is geographic, ecological or reproductive. In the 
following account the groups concerned are defined first, and thus the descriptions 
of species offered are concise, depending more on evidence of species isolation 
than indefinite morphological differences. 


(a) “* exigua’’ group 

Small species of Patiriella in which F rarely exceeds 15mm. The number 
of rays is usually five and actinal intermediate spines are usually single. 
Abactinal spines from the centre of the disc are from 0-2 to 0:3 mm. in length. 


Patiriella exigua (Lamarck, 1816) 
(Plate Iv (c)) 
Restricted synonymy: Asterias exigua Lamarck, 1816; Asterina krausw Gray; 
1840; Asteriscus pentagonus Miller and Troschel, 1842 (in part); Asterina 
exigua Perrier, 1876; Patiriella exigua Verrill, 1913 (in part); Asterina exigua 
Mortensen, 1921; Patiriella eaigua H. L. Clark, 1938, 1946 (in part) ; HEndean, 
1956 (in part); Shepherd, 1968; Dartnall, 1970b. 

Non: Asteriscus pentagonus Miller and Troschel, 1842 (in part); Asterina 
exigua H. L. Clark, 1908; Koehler, 1910; Fisher, 1919; H. L. Clark, 1921 ; 
Patiriella exigua Livingstone, 1932; Asterina (Patiriella) exigua Mortensen, 
1933a; Domantay and Roxas, 1938; Engel, 1938. 

Asterina minuta of various authors has not been included in the synonymy 
because of lack of precise information about localities. However, Gray (1840) 
was clear that the specimens he attributed to A. minuta were from the ‘ West 
Indies, St. Vincents”’. If those details are correct then those specimens are 
probably Liitken’s Asterina foliwm or a related form. Information about A. 
folium with which to compare Gray’s A. minuta is available in a paper by de Roa 
(1967). As far as synonymy is concerned the complex of asterinids from South 
Africa has also been ignored in this paper as it is commented upon by both H. L. 
Clark (1923) and Mortensen (1933a). 


Asterias exigua Lamarck was described from material whose provenance 
was suggested to be the Americas (Perrier, 1876). Perrier examined Lamarck’s 
seven type specimens and also noted that a further series of nine specimens, 
collected from the Cape of Good Hope in 1829 by M. Raynaud, were labelled 
Asterias exigua by Lamarck. Perrier (loc. cit.) and Koehler (1910) both 
recognized Asterias exigua as a valid species containing the synonyms Asterias 
minuta de Blainville, 1834; Agassiz, 1834 (in Agassiz, 1838) and Gray, 1840 ; 
Asteriscus pentagonus Miiller and Troschel, 1842 and Asterina pentagona von 
Martens, 1866. 


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A. J. DARTNALL 41 


Patiriella exigua is recorded in the literature from throughout the Australo- 
Indo-west-Pacific from South Africa to the Philippines, the New Hebrides and 
the eastern and southern coasts of Australia including Tasmania. Whitelegge 
(1889) was the first to note the orally directed gonoducts of the adult and the 
shortened development of the larvae. Mortensen (1921) confirmed Whitelegge’s 
observations and suggested that P. erigua might contain more than one species, 
quoting Ludwig that Asterina pentagona, an accepted synonym, possessed 
genital pores opening on the abactinal surface. Since that time abbreviated 
development has been accepted as characteristic of the species and an account 
is incorporated in at least one standard textbook of zoology (Parker and Haswell, 
1962). 


In September, 1967, Dr. J. R. Grindley obtained for me material attributed 
to Patiriella exigua from Port Elizabeth, South Africa. This material can be 
divided into two groups: 

(1) Asterinid sea stars with abactinally directed gonoducts, attaining 18 mm. 
R in these samples and whose actinal surface is not consistently blue-green in 
colour. Miss A. M. Clark of the British Museum (Natural History) was kind 
enough to compare some of these specimens with material in her care and informs 
me that they agree best with Asterina dyscrita H. L. Clark, 1923. 


(2) Asterinids with actinally directed gonoducts, attaining a maximum R 
of approximately 12mm. and whose actinal surface is always blue-green in 
colour. 


The latter group is identical to that species of sea star from south-eastern 
Australia commonly determined as Patiriella exigua and it was decided to 
investigate that species throughout its known range. Through the co-operation 
of many individuals and institutions, material was obtained from areas throughout 
most of the recorded distribution of the species and all the specimens were 
examined to determine the position of the gonopores and the orientation of the 
gonoducts. Morphological differences between the samples were slight and 
gonopore position remained the one constant character for investigation in 
preserved material. The map (Fig. 1) shows the localities from which material 
was obtained. Orientation of gonoducts in the samples is indicated by the 
arrowheads and the following distribution is clear. 


North of about 28°S. in Australia all the samples attributed to Patiriella 
exigua possessed abactinally orientated gonopores. 


In Australia the form with actinally directed gonopores is found along the 
east coast, south of approximately 28°S., to Port Lincoln on the south coast 
at approximately 136° E. The range of this form extends southwards to 
encompass the coasts of Tasmania and eastward of the continent to Lord Howe 
Island at about 159° EK. This form is also found in South Africa and the isolated 
islands of St. Paul and New Amsterdam. 


Application to Dr. G. Cherbonnier revealed that Lamarck’s types of Asterias 
exigua are no longer in Paris. Dr. E. Binder of Geneva and Dr. W. Vervoort 
of Leiden inform me that neither of their institutions hold Lamarckian echinoderm 
types. Further enquiries suggest that Lamarck’s specimens no longer exist 
and in order to preserve the name “‘ exigua”’ for this well-documented animal it 
appears necessary to erect a neotype for the species. 


The Cape of Good Hope has been accepted as the type locality by various 
authors, but without a locality for Lamarck’s specimens this is very uncertain. 
Among those specimens available to me the two from the Cape do not demonstrate 
adequately the key character to the species, namely the gonopores, so I have 
chosen a recent specimen from False Bay, South Africa, which lies to the east 
of the Cape of Good Hope and is now designated the type locality. 


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42 SEA STARS GENUS PATIRIELLA 


Details of neotype: T.M. H508. R=11-5mm., from the type locality, False 
Bay, South Africa (34°8./18° E.); 10.x.1969; don. J. G. Field. 

Other material examined : 

A.M. J7762 (6), City of Melbourne Bay, King Is., Tasmania, June 1968, 


S. Kerrison. 


B.M. 1840.11.25.1 and 1840.11.25.5, Cape of Good Hope. Both specimens © 


are syntypes of Asterina kraussu Gray. 

M.N. 6682 (35), St. Paul Island, 26.iv.1903, Deutche Stdpolar-Expedition 
1901-1903 ; 6683 (6), Amsterdam Is., 27.iv.1903, Deutche Stidpolar-Expedition. 
N.M.V. H151 (6), Flinders, Victoria, 6.vi.1969, A. J. and P. J. Dartnall. 

Q.M. G3765 (4), Currumbin, Queensland, 8.vili.1953, R. Endean. 


obscura 


seudoexigua 
iy p Gg 


_exigua 


brevispina 


regularis ‘\ 
Fig. 1. Contemporary distribution of the species of Patiriella found in Australia. P. nornata 
from Western Australia is omitted. 


Inset: Map showing the localities from which material of the “ exigua’’ group has been 
obtained. Orientation of gonopores is shown by the solid triangles— 4 abactinal; © actinal. 


“ 


T.M. H212 (2), Roaring Beach, Port Davey, Tas., Feb. 1937, C. Davies ; 
H222 (12), Stanley, Tas., Oct. 1937, A. W. G. Powell; H332 (31), Nubeena, 
Tas., 14.v.1967, L. Jenkins ; H345 (2), Murdunna, Tas., 12.1.1967, A. J. Dartnall ; 
H349 (8), Port Arthur, Tas., 20.iv.1967, A. P. Andrews and EH. Aves ; H365 (12), 
Granville Harbour, Tas., 6.xi.1967, A. J. Dartnall ; H392 (6), Dennes Pt., Bruny 
Is., Tas., 24.1.1968, A. J. Dartnall; H409 (13), South Australia, 1967, S. A. 
Shepherd ; H452 (4), Maria Is., Tas., 15.iv.1968, A. J. Dartnall; H456 (4), 
Swan Is., Tas., Sept. 1968, F. L. Sutherland ; H458 (9), Cape Portland, Tas., 
2.11.1969, A. J. Dartnall; H479 (18), Salmon Reef, Lord Howe Is., 6.xi.1968, 
W. G. Thornton ; H480 (6), Trouser’s Point, Flinders Is., Tas., 7.1.1969, T. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wates, Vou. 96, Part 1 


A. J. DARTNALL 43 


Goede ; H483 (7), Marrawah, Tas., 1.xii.1968, B. J. Smith; H484 (5), Greens 
Beach, north Tas., 13.viii.1969, RK. H. Green; H485 (10), Stevens Bay, Port 
Davey, Tas., 8.1.1969, D. Milledge ; H486 (10), Long Reef, near Sydney, N.S.W., 
1.vii.1969, E. C. Pope; H496 (5), Salt Creek, Port Lincoln, S. Australia, 
10.xi1.1969, J. Veitch; H494 (6), Port Elizabeth, South Africa, Sept. 1967, 
J. R. Grindley ; H498 (8), False Bay, South Africa, 34°S./18° E., 10.x.1969, 
ay Gap ield. 

UNIVERSITY OF CAPE TOWN: ECOLOGICAL SURVEY COLLECTIONS. All from 
South Africa. L10 (8), East London, 32°8./28° E., 6.vii.1937; LB112 (4), 
Langebaan Lagoon, 33°8./18° H., 15.vii.1946; N35 (6), Port Nolloth, 
29°S./16° H., 27.x.1935; LU3A (13), Luderitz, 26°S./15° E., 15.vii.1946; 
D24 (8), Durban, 29°8./15° E., 27.vi.1935 ; AT (10), Oudekraal, 34°8./18° E., 
2.11934; E3 (9), Port Elizabeth, 33°8./28° H., 3.vii.1936 ; S9 (10), Still Bay, 
34°S./21° K:, 41,1932. 

w.A.M. 37-69 (1), Sydney, N.S.W., late 1963, C.S.I.R.O. Fisheries ; 

39-69 (3), Gunnamatta Bay, Port Hacking, N.S.W., 31.x.1963, C.S.I.R.O. 
Fisheries. 
Description: As above, in the description of “ exigua” group. Oral spines 5-6 ; 
suboral spines 1; furrow spines 2 to each adambulacral ossicle and basally 
webbed ; subambulacral spines 1. Actinal intermediate spines 1, 2 towards 
the dise margin, absent proximal to the mouth plates. In preserved specimens 
the plate at the apex of the chevron which defines the actinal intermediate 
area may “‘ float ”’ clear of its fellows on a sheet of membrane. A pair of gonopores 
are present in each actinal intermediate area about level with the third ambulacral 
ossicle and between the second and third chevrons of actinal intermediate plates 
(Plate IV, c). 


R:r about 1:2-1-3:1. 


Diagnosis: A small species of Patiriella with actinally directed gonoducts and 
abbreviated larval development. 

Colour in life: The actinal surface is consistently blue-green. The ground colour 
of the abactinal surface is dull green or brown. Red, orange, purple, brown or 
pastel greens or browns on the spines grouped at the inferomarginal fringe, along 
the top of the rays and at the disc centre, often produce an attractive geometric 
pattern against the ground colour. The tissues of the circum-oral ring are 
blue-green. The tube feet are blue-green and the suckers off-white. The 
papulae are transparent, showing blue-green coloration at their exit from the 
papular pores. 

Distribution: South Africa, St. Paul group, Australia south of 28°S. and E. of 
136° E., Tasmania, Lord Howe Island. 

Habitat : Mid-tide level on rocky shores. In areas where wave splash is consistent 
the species may be found high on the shore. Specimens from South Australia 
were taken under rocks on a muddy substrate and the species has been collected 
on sand at Anson’s Bay, N.E. Tasmania. In Australia Patiriella exigua is often 
associated with the alga Hormosira banksi. 


Patiriella pseudoexigua sp. nov. 

(Plate Iv (a)) 
Synonymy: % Asteriscus pentagonus Miller and Troschel, 1842 (in part), 
2? Asterina pentagona von Martens, 1866; Asterina exigua H. L. Clark, 1908 ; 
? Koehler, 1910; Patiriella exigua Verrill, 1913 (in part); Asterina exigua 
Fisher, 1919; Asterina exigua H. L. Clark, 1921 ; Patiriella exiqua H. L. Clark, 
1938 (in part); 2? Asterina (Patiriella) exigua Domantay and Roxas, 1938 ; 
Engel, 1938; Patiriella exigua H. L. Clark, 1946 (in part); Endean, 1956 (in 
part). 


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44 SEA STARS GENUS PATIRIELLA 


Ludwig (in Mortensen, 1921) observed that Asterina pentagona von Martens 
possessed genital pores opening on the abactinal surface of the animal in contrast 
to Patiriella exigua. Thus it would appear that the name pentagonus of Miller 
and Troschel is available to accommodate the species of Patiriella found north 
of 28°S. in Australia. However, two species of Patiriella are present in north- 
eastern Australia and I find it impossible to decide which is actually pentagonus 
of past authors. Accordingly both this species and one belonging to the 
‘“‘ regularis > group are described as new. Future workers in the Indo-west- 
Pacific may need to place one of these species in synonymy if further information 
validates the status of Miller and Troschel’s species name. The alternative 
is to declare formally that pentagonus is invalid, but this must await further 
studies on the species of Patiriella in the waters north of Australia. 


Holotype: T.M. H499. A dried specimen, K=12-5mm., Airlies Beach, 
Proserpine, Queensland (type locality), 12.iv.1970, A. J. Dartnall. 

Paratypes: T.M. H500 (6), Airlies Beach, Proserpine, Queensland, 12.iv.1970, 
A. J. Dartnall; A.M. J7761 (2), Airlies Beach, Queensland, September/October 
1968, K. Deacon. 


Other material examined : 

B.M. 1890.5.7.581-584 (4), Zamboanga, Philippines; 1968.6.14.156-160 
(5), Solomon Islands. 

M.C.Z. 2297 (1), Murray Is., Mer., Oct. 1913, H. L. Clark, Carnegie 
Expedition ; 2083 (15), Espiritu Santo, Malo, New Hebrides, January 1911, 
Rev. J. Annand; 2296 (23), Erub, Torres Strait, 19.ix.1913, H. L. Clark ; 
2298 (1), Hrub, Torres Strait, 19.ix.1913, H. L. Clark. 

N.M.vV. H116 (1), Aola, Guadaleanal, 1901, G. Officer; G114 (1), Bora 
Bada, New Guinea, March 1891, J. Exton; H110 (1), Bingal Bay, Queensland, 
July 1964, J. Kerslake. 

Q.M. G38767 (2), Port Curtis, Queensland, 16.vili.1961, R. Endean ; G3762 
(26), South Yeppoon, Queensland, 15.1.1953, R. Endean ; G3764 (22), Airlies 
Landing, Proserpine, Queensland, 21.iv.1953, R. Endean. 

T.M. H501 (32), Airlies Beach, Proserpine, Queensland, 12.iv.1970, A. J. 
Dartnall; H502 (4), one mile N.E. Airlies Beach, Proserpine, Queensland, 
12.iv.1970, A. J. Dartnall; H503 (6), south end of King’s Beach, Bowen, 
Queensland, 13.iv.1970, A. J. Dartnall. 

U.S.N.M. 38088 (12), Philippine Islands, E. A. Mearns; 40261 (7), Port 
Binang, Subie Bay, Philippines, 8.i1.1908, Albatross Philippines Expedition 
1907-1910 ; 40254 (4), Borneo, 2.i11.1908, Albatross Expedition. 

W.A.M. 43-69 (3), Real Point, Villa Carmen, Cabcaben, Bataan, Philippines, 
24.vii.1965, Western Australian Museum Luzon Expedition 1965. 


Description of holotype: A small asterinid sea star very similar in morphology 
to Patiriella exigua. R:r about 1-3:1. Rays five. Each oral plate with five 
oral spines and one suboral spine. Most furrow spines are paired except near 
the mouth plates, where two or three adambulacral plates carry three furrow 
spines. The subambulacral and actinal intermediate spines are each placed 
singly to an actinal plate. There are no pairs of spines towards the edge of the 
dise and spines are absent near to the mouth plates. The inferomarginal plates 
carry seven to nine spines ; the abactinal plates four to 20 spinelets. 


The gonoducts are orientated abactinally and the gonopores open on the 
abactinal surface. 
Diagnosis: A small species of Patiriella with abactinally directed gonoducts. 


Colour in life: The actinal surface is a dull brownish-green; the abactinal 
surface dull brown and green. Most specimens are darker on the centre of the 
dise and along the crest of the rays, forming a five-rayed star pattern on the 


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A. J. DARTNALL A5 


abactinal surface. Variants show some russet and orange patches in the inter- 
radial areas. The circum-oral ring is bluish, the tube feet pale straw coloured, 
and the suckers off-white. 


Distribution: The east coast of Australia north of about 28°S., Torres Strait, 
New Guinea, the New Hebrides, the Philippines and Borneo. 


Habitat: Under rocks at mid-tide level. 


Patiriella vivipara Dartnall, 1969 
Diagnosis: A small species of Patiriella with no gonoducts ; coelomic incubation 
of young; viviparous. 
Distribution: Restricted to S.E. Tasmania. 


(b) “‘ regularis”’ growp 

Species of Patiriella resembling the ‘ exigua”’? group morphologically, in 
which F& exceeds 15mm. and the abactinal spinelets are longer than in the 
““ exigua’’ group (between 0-4 and 0-9 mm. in length). 


Patiriella regularis (Verrill, 1867) 
Restricted synonomy: Asterina (Asteriscus) regularis Verrill, 1867; Patiriella 


regularis Verrill, 1913 ; Patiriella mimica Livingstone, 1933 ; Patiriella regularis 
Dartnall, 1970. 


Diagnosis: A species of Patiriella with five rays. RK: r=1-5-1-8:1. 
Distribution: New Zealand and 8.K. Tasmania. 


Patiriella inornata Livingstone, 1933 


Diagnosis: A species of Patiriella with five rays, short furrow spines and the 
innermost oral spine notched at the tip. 


Distribution: Western Australia. 
Remarks: This species is known only from the holotype. 


Patiriella nigra H. LL. Clark, 1938 
Diagnosis: Five rays. Actinal intermediate area with dense spinulation ; 
many plates towards the inferomarginal fringe carrying two or three spines. 
On the abactinal surface the spinulation of the papulate areas is distinct, 
approaching the condition described for species of Paranepanthia. Colour in 
life black. The gonopores open on the abactinal surface. 
Distribution: Lord Howe Island. 


Patiriella obscura sp. nov. 
(Plate Iv (b)) 
Synonymy: 7 Patiriella exgua Livingstone, 1932; Patiriella exigua Endean, 
1956 (in part). 

Museum samples showed that a large species of Patiriella was present in 
north-eastern Australia and probably extended to the Philippines. When I first 
collected this animal alive I considered it to be the adult of Patiriella pseudo- 
exigua sp. nov. However, examination showed that the gonads of P. obscura 
were not mature until 13mm. Rk was attained and P. pseudoexigua possesses 
mature gonads at 11 mm. R. 

Holotype: T.M. H504. A dried specimen, R=18mm., Rose Bay, Bowen, 
Queensland (type locality), 13.iv.1970, A. J. Dartnall. 


Paratypes: T.M. H505 (6) and A.M. J7791 (6), Rose Bay, Bowen, Queensland, 
13.iv.1970, A. J. Dartnall. 


Other material examined : 


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46 SEA STARS GENUS PATIRIELLA 


Q.M. G5330 (2), Rowe’s Bay, Townsville, Queensland, August 1967, R. 
Monroe; G3763 (5), Low Island, Queensland, 13.viii.1954, R. Endean. 


T.M. H506 (15), Rose Bay, Bowen, Queensland, 13.iv.1970, A. J. Dartnall ; 
H507 (23), Rowe’s Bay, Townsville, Queensland, 14.iv.1970, A. J. Dartnaill. 


COLL. ZOO. DEPT., UNIVERSITY OF THE PHILIPPINES. Two dried specimens 
from the vicinity of Port Galera Bay. 


Description of holotype: A species of Patiriella with five rays. R:r=1-2:1. 
Each oral plate carries five or six oral spines and one or two suboral spines. 
Most furrow spines are arranged in pairs, but two adambulacral plates near the 
mouth carry three spines. Subambulacral spines are single with occasional 
pairs. Actinal intermediate spines are mainly single with occasional pairs near 
the edge of the disc. Fringing spinelets are in groups of seven to 13. The 
abactinal spinelets range from four to 20 to an abactinal plate. 


The gonopores open on the abactinal surface. 


Diagnosis: A species of Patiriella of R >15mm. with abactinally directed 
gonoducts. Suboral spines often paired. 


Colour in life: The actinal surface is dull brownish-green ; the abactinal surface 
dull brown, reddish and purple. Dull, pale colours show indistinct geometric 
patterns against the ground colour. Tube feet greenish-blue ; suckers off-white. 
Circum-oral ring bluish. 


Distribution: N.E. Australia to the Philippines. 


Habitat: Under rocks at mid-tide level. This species will feed over the surface 
of mud surrounding rocks at low tide and the body colour provides very effective 
camouflage at this time. 


Patiriella calear (Lamarck, 1816) 
Restricted synonymy: Asterias calear Lamarck, 1816; Asterina calear McCoy, 
1890; Patiriella calear Verrill, 1913; H. L. Clark, 1946; Endean, 1953 ; 
Shepherd, 1968; Dartnall, 1970b. 


Diagnosis: A species of Patiriella with seven to 11 rays, usually eight. Apart 
from the number of rays this species falls most naturally into the “ regularis ” 
group. 

Distribution : From the south coast of Western Australia to southern Queensland 
and the coasts of Tasmania. 


Remarks: It appears that Lamarck’s types of Asterias calcar no longer exist. 
I have not erected a neotype for this species because I am not aware that the 
identity of this well marked species has been questioned. 


(c) “ gunni”’ group 
Species of Patiriella with six rays and the spines of the actinal intermediate 


area arranged in pairs. Abactinal spines near centre of the disc between 0-2 
and 0-4 mm. in length. 


Patiriella gunnu (Gray, 1840) 


Restricted synonymy: Asterina gunnii Gray, 1840; Patiriella gunnii Verrill, 
1913; H. L. Clark, 1946; A. M. Clark, 1966; Shepherd, 1968; Dartnall, 
1970b. 

Diagnosis: A flattened form with paired subambulacral spines. Maximum 
R about 39 mm. 


Distribution: Known from Tasmania, Western Australia to New South Wales, 
Lord Howe Island. 


PROCEEDINGS OF THE LINNEAN SocrtEty or NEw SoutH Watss, Von. 96, Part 1 


A. J. DARTNALL ATT 


Patiriella brevispina H. L. Clark, 1938 


Diagnosis: An arched form with single subambulacral spines attaining about 
52mm. R. 


Distribution: Western Australia to New South Wales, the north coast of 
Tasmania. 


Key to the Australian Species of Patiriella 
I Actmal intermediate spmes paired; rays six......................-- ~“ gunnit”’ group (2) 
Actinal intermediate spines single ; rays usually five or more than six.................- (3) 
2. (1) Flattened ; subambulacral spines usually paired ; Colour in life varied... .Patiriella gunnii 
Arched; subambulacral spines usually single; colour consistently and uniformly 
DULL Ops tse oito cs sepew nue) sens crisis, oc ua eat oon aU ley ANE cage Mrs Ceaaeel sae Patiriella brevispina* 
3. (1) Small; R<15mm., actinal spines <0-4mm. in length “ exigua’’ group (4). 
Large; R>15mm., actual spines >9-4 mm. in length “ regularis ” group (6). 


4. (3) Gonoduct absent ; coelomic incubation of young.................. Patiriella vivipara 
Conoductiypreseiibry suteel else sts, caine Areas ee eee teean ores iovace MERRIE: ba scee AA APRS (5) 
5. (4) Gonoducts directed actinally. Actinal surface blue-green.............. Patiriella exigua 
Gonoducts directed abactinally. Actinal surface dull brownish green Patiriella pseudoexigua 
G3). ERS TS cia vetics fs o:8 sta Dra cnet Si ars eee nen Grail nicl Ue a AT Ie, Goa kh o eae Patiriella calcar 
TRONS. Wick date ane per eae ee ELS CEM Mee ig ofa ret ten teen ee Om Ses 4. ol ca he aera eee (7) 
7 (6) Spinulation of apapulate and papulate abactinal areas distinct; colour black (Lord 
HEVOWOpUSH) teeter ee Cee ee ee els ences ea RO Ee ne EO oie Patiriella nigra 
Spinulation of abactinal interradial areas not as dense as above ; colour not black.. . .(8) 
8. (7) Innermost oral spine flat, very wide notched at tip.................... Patiriella inornata 
ianermosthoralespine NOt sasealboOven- eee soe Don ae eo ee aera a acre: (9 
9. (8) Suboral spines single; colour of actinal surface off-white............ Patiriella regularis 


Suboral spines often paired ; colour of actinal surface dull brownish-green Patiriella obscura 


* In a previous key to the Asterinidae of Tasmania (Proc. Roy. Soc. Tas., 1970) this section 
runs down to P. calcar. The opportunity is taken to rectify this mistake here. 


DISCUSSION 


The distribution of species within the “‘ exigua’”’ group shows a sequence in 
which each species is separated from the next by a reproductive mechanism. 
These observations reinforce the concept of sibling pairs of species mooted in a 
previous paper (Dartnall, 1970b) except that a triple sequence is now discernible. 
Patiriella pseudoexigua, with upwardly directed gonoducts and presumably free- 
swimming larvae, is found north of 28°S. in eastern Australia. South of 28°S. 
and isolated from P. pseudoexigua by a reproductive ‘‘ inversion ”’ is Patiriella 
exigua. The third species in the sequence is the hermaphrodite, viviparous form, 
Patiriella vivipara, which is restricted to the cool coasts of S.E. Tasmania. 


The discovery of a “ regularis’ form (Patiriella obscura) in N.K. Australia 
adds weight to the concept that the precursors of Patiriella came from the Indo- 
west-Pacific area. It is suggested that the ‘‘ regularis ’’ forms have contributed 
to the New Zealand fauna (Patiriella regularis) by way of the Lord Howe Rise 
(Patiriella nigra) and to south-eastern Australia through evolution of Patiriella 
calear. It is possible that the “ exigua’”’ forms represent an older invasion as 
the contemporary distribution of P. exigua encompasses the temperate and cool 
temperate waters of both Australia and South Africa and the intervening St. Paul 
group of islands (see Fig. 1). 


Figure 1 shows the present distribution of the Australian species of Patiriella. 
The limits of distribution of P. exigua and P. pseudoexigua at about 28° S. roughly 
correspond with the biogeographic boundary defined amongst others by Endean 
(1957) when he considered the biogeography of the shallow water echinoderm 
fauna of Queensland. When the distribution of the species belonging to the 
“ gunnit” group is also considered it can be seen that the agreement with the 
marine zoogeographic provinces of Bennett and Pope (1953, 1956) is marked 
and that the distributional limits accord well with those postulated by George 
(1969) for the tropical and subtropical Palinuridae. 


PROCEEDINGS OF THE LINNEAN Socrety oF NEw SoutH WALES, VoL. 90, Part 1 


48 SEA STARS GENUS PATIRIELLA 


In an earlier paper (Dartnall, 1970a) the distributions of the hermaphrodite 
forms known within the Asterinidae were shown to lie in widely separate areas 
around 40° of latitude, both north and south. This evidence must now be 
restricted to the non-fissiparous asterinids because Dr. R. Kenny has sent me 
samples of hermaphrodite individuals of Nepanthia belcheri (Perrier) from 
Moreton Bay, Queensland. When the distribution of the species belonging to 
the ‘*‘ exigua ”’ sequence is considered, the speculations which follow are of interest. 

(a) The distributions of the ‘‘ exigua”’ species are defined by thermal 
tolerance alone, the observed sequence of reproductive mechanisms being 
fortuitous. 

(b) Each kind of reproductive mechanism is limited by temperature, higher 
latitudes and lower temperatures directing the success of hermaphrodite forms, 
species with abbreviated development, or a combination of both. 


It is probably not possible to test these speculations. However, it may be 
possible to find out whether similar reproductive sequences occur in other groups 
of Asteroidea along the long eastern seaboard of Australia. 


ACKNOWLEDGEMENTS 


I would thank the following persons and the institutions that they represent 
for help with material and literature: Dr. B. Z. Anicete, University of the 
Philippines ; Dr. B. Campbell, Queensland Museum; Miss A. M. Clark, British 
Museum (Natural History); Miss M. E. Downey, United States National 
Museum; Professor H. B. Fell, Museum of Comparative Zoology, Harvard ; 
Mr. J. G. Field, University of Cape Town; Dr. J. R. Grindley, Museum, Snake 
Park and Oceanarium, Port Elizabeth, South Africa ; Dr. R. Kenny, James Cook 
University of North Queensland; Dr. and Frau D. Kithlman, Humboldt 
University of Berlin; Miss E. C. Pope, Australian Museum ; Mrs. 8. Slack-Smith, 
Western Australian Museum ; and Dr. B. J. Smith, National Museum of Victoria, 


I am grateful to the Trustees of the Tasmanian Museum and Art Gallery, 
who granted leave of absence to study species of Patiriella in Queensland, and 
the Trustees of the Science and Industry Endowment Fund, ©.S.I.R.0O., who 
made a grant-in-aid of this study. 


Finally, grateful acknowledgement is made to Dr. W. Bryden, Dr. E. R. 
Guiler, Dr. D. Martin and Sir Henry Somerset, who have freely offered advice 
and encouragement at various stages of this work. 


References 


Acassiz, L., 1838.—Prodromus of a Monograph of the Radiata and Echinodermata III. Ann. 
Mag. Nat. Hist., 1: 440-449. 

BENNETT, I., and Popr, EH. C., 1953.—Intertidal Zonation on the Exposed Rocky Shores of 
Victoria, together with a rearrangement of the biogeographical provinces of temperate shores. 
Aust. J. Mar. Freshw. Res., 4: 105. 

, 1960.— Intertidal Zonation of the Exposed Rocky Shores of Tasmania and its Relation- 
ship with the Rest of Australia. Aust. J. Mar. Freshw. Res., 11: 182. 
Buainvitte, H. M. D. de, 1834.—Manuel d’actinologie. ou de zoophytologie, 688 pp. 
Brine A. M., 1966.—The Port Philip Survey 1957-63. Echinodermata. Mem. Nat. Mus. Vic., 
7: 289-384. 
CrarKk, H. L., 1908.—Echinodermata. Sci. Res. ‘“‘ Thetis’. Mem. Aust. Mus., 4: 519-564. 
, 1921.—The Echinoderm Fauna of Torres Strait. Pap. Tortugas Lab., 10: 1-244. 
, 1923.—The Echinoderm fauna of South Africa. Ann. S. Afr. Mus., 13: 221-428. 
, 1938.—Kchinoderms from Australia. Mem. Mus. Comp. Zool. Harvard, 55: 1-596. 
, 1946.—The Echinoderm fauna of Australia. Publ. Carneg. Instn., 566: 1-567. 

Darrnat., A. J., 1969a.— New Zealand Sea Stars in Tasmania. Proc. Roy. Soc. Tas., 103: 53-55. 

, 1969b.—A viviparous species of Patiriella (Asteroidea, Asterinidae) from Tasmania. 

Proc. Linn. Soc. N.S.W., 93 (2): 294-296. 

, 1970a.—Some species of Asterina from Flinders, Victoria. Vict. Nat., 87 (1): 1-4. 
, 1970b.—The Asterinid Sea Stars of Tasmania. Proc. Roy. Soc. Tas., 104: 73-77. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Watss, Vou. 96, Part 1 


PLATE It 


Proc. Linn. Soc. N.S.W., Vol. 96, Part 1 


‘wu Q| 


1 
i 
j 1 le) Ok | eae 
OPM Oe ke re) eh ha, Py 
} ; ‘i , oN ie Co wsTih (wl ; 
pena 
2 AL ee 
: re ] el We Re 4 oy 
a ee, 9 als ry oy 
‘ ak , ; aN hee ya Vi iu 
, : : fale si ate EM 
" Lie + i - : ¥ _* 


, Mf pin aia Nay 


Proc. Linn. Soc. N.S8.W., Vol. 96, Part 1 PLATE IV 


A. J. DARTNALL 49 


Domantay, J. 8., and Roxas, H. A., 1938.—The littoral Asteroidea of Port Galera Bay. Philip- 
pine Jour. Scv., 65: 203-237. 

ENDEAN, R., 1952.—Queensland faunistic records. Part III. Echinodermata (excluding 
Crinoidae). Pap. Dep. Zool. Univ. Qid., 1 (3): 53-60. 

, 1956.—Queensland faunistic records. IV. Further records of Echinodermata 
(excluding Crinoidea). Pap. Dep. Zool. Univ. Qid., 1 (5): 121-140. 

, 1957.—The biogeography of Queensland’s shallow-water echinoderm fauna (excluding 
Crinoidea), with a rearrangement of the faunistic provinces of tropical Australia. Aust. J. 
Mar. Freshw. Res., 8 (3): 233-273. 

ENGEL, H., 1938.—Resultats Scientifiques du Voyage aua Indies Orientales Neerlandaises, 
III (1)8. 

FisHer, W. K., 1919.—Starfishes of the Philippine Seas and adjacent waters. Bull. U.S. Nat. 
Mus., 100 (3): 1-711. 

GrorcE, R. W., 1969.—Natural distribution and speciation of marine animals. J. Proc. Roy. 
Soc. West. Aust., 52 (2): 33-40. 

Gray, J. E., 1840.—A synopsis of the genera and species of the class Hypostoma (Asterias 
Linneus). Ann. Mag. Nat. Hist., 6 (1): 175-184, 275-290. 

KoEHLER, R., 1910.—The Shallow Water Asteroidea. Hchinoderma of the Indian Museum, 
6: 1-191. 

LAMARCK, J. B. P., 18146. — Histoire naturelle des animaux sans vertebres, 2: 522-568. 

Livinestonge, A. A., 1932.— Sci. rept. Great Barrier Reef Exped. 1928-29, 1V (8): 241-265. 

, 1933.—Some genera and species of the Asterinidae. Rec. Aust. Mus. Sydney, 19: 1 -20. 

Martens, E. von, 1866.—Uber ostasiatische Echinodermen. Arch. f. Naturgesch, 32: 57-88, 
133-189. 

McCoy, F., 1890.—Prodromus of the Zoclogy of Victoria, 2 (20): 371-375. 

Mortensen, T., 1921.—Studies of the development and larval forms of echinoderms, Copenhagen 
C.E.G.: 183-190. 

, 1921.—Echinoderms of South Africa (Asteroidea and Ophiuroidea). Vidensk. Medd. 

naturh. Foren. Kbh., 93: 215-400. 

MUuer, J., and TrRoscHEt, F. H., 1842.—System der Asteriden, xx +135. 

Parker, T. J., and HASweELi, W. A., 1962.—Textbook of Zoology, 1: 1-700. McMillan, London 
(6th ed.). 

PERRIER, E., 1876.—Revision de la collection de stellérides du Muséum. Arch. zool. exp., 5: 
1-104, 209-304. 

Roa, E. Z. de, 1967.—Contribuction al estudio de los equinodermos de Venezuela. Acta Biologica 
Venezuelica, 5 (17): 267-333. 

SHEPHERD, S. A., 1968.—The Shallow Water Echinoderm Fauna of South Australia. I. The 
Asteroids. Rec. S. Aust. Mus., 15 (4): 729-756. 

VERRILL, A. E., 1867.—Notes on the Radiata in the Museum of Yale College. Trans. Conn. 
Acad. Art. Sci., 35 (4): 247-351. 

, 1913.—Revision of the genera of starfishes of the subfamily Asterinidae. Amer. 
Jour. Sci., 35 (4): 477-485. 

WHITELEGGE, T., 1889.—Invertebrate fauna of Port Jackson. Subkingdom Echinodermata. 
Proc. Roy. Soc. N.S.W., 23: 197-206. 


EXPLANATION OF PLATES 
PLATE IIt 


(a) Patiriella pseudoexigua sp. nov. Actinal surface of holotype (T.M. H499). 
(6) Patiriella obscura sp. nov. Actinal surface of holotype (T.M. H 504). 


PLATE IV 
(a) Airlie’s Beach, Proserpine, Queensland. Type locality of P. pseudoexigua sp. nov. 
(b) Rose Bay, Bowen, Queensland. Type locality of P. obscura sp. nov. 
( 


c) Patiriella exigua (Lamarck). Actinal surface of neotype (T.M. H499). The positions of the 
gonopores in two interradii are indicated by the arrowheads. 


PROCEEDINGS OF THE LINNEAN Society or NEw SourH WaAtgEs, Vou. 96, Part 1 


A SEA STAR OF GENUS CTENODISCUS FROM TASMANTA 


A. J. DARTNALL 
The Tasmanian Museum, Hobart, Tasmania 


[Accepted for publication 23rd September 1970] 


Synopsis 
A single specimen from deep water off northern Tasmania constitutes the first record of the 
sea star family Goniopectinidae from Australian waters. 


INTRODUCTION 


The sea star genus Ctenodiscus of the family Goniopectinidae is represented 
by two species in the Southern Hemisphere. Ctenodiscus australis Liitken 
is known from waters off the east coast of South America and C. procurator 


Hh NOB 
BB CUT pt 
uy 


11))) 1) 2 


) WT) 1 5 


YS vi 


1 
UN PM 


TUN LN)! VN) 


Fig. 1. Ctenodiscus sp. Side view from tip of ray to midline of the interradius. 


Sladen from the west (Bernasconi, 1964). Ctenodiscus orientalis Fisher is tropical 
Indo-Malay in distribution and C. crispatus (Retzius) is an arctic-boreal form 
with a circumpolar distribution (D’yakonov, 1950). 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wass, Vou. 96, Part 1 


A. J. DARTNALL 51 


The single specimen discussed here was taken on the 24th September, 1953, 
from the F.R.V. Derwent Hunter 45 miles H.N.E. of Stokes Point, King Island, 
in 300-240 fathoms (546-436 metres). I am grateful to Mr. A. M. Olsen, who 
collected the specimen, which is now housed in the collections of the Tasmanian 
Museum. 

Ctenodiscus sp. (ef. C. orientalis Fisher) 

One dried specimen. Tasmanian Museum Reg. No. H497. R=19-5mm., 
ry=8-0 mm., br at base of ray=7-:0 mm., &: 7 2-43:1. 

Description: A sea star with five pointed rays and deep interbrachial arcs. The 
aboral surface is flattened and the oral surface is inclined at a steep angle, the 
mouthplates being the lowest point (Fig. 1). 

The actinal surface is covered with small paxillae that have no central 
spinelets and four to eight peripheral spinelets. Near the dise centre a small 
area (about 2-5 mm. across) is more closely covered with paxillae. This may be 
a retracted epiproctal cone. 

The madreporite is some 2 mm. across and placed close to the margin. The 
furrows of the madreporite are roughly parallel and aligned along the radial 
axis of the body in which the madreporite lies. 

The body is bordered by two rows of marginal plates forming an upright wall. 
Opposite inferomarginal and superomarginal plates are approximately equal in 
height, there being 16 marginals along the side of each ray. ‘The largest pairs of 
inferomarginal and superomarginal plates are the third, fourth and fifth from the 
interradial line, the height of the marginal wall being reduced from those plates 
both to the tip of the ray and the mid-line of the interradius. Fascioles are 
present between all the marginal plates. The fasciolar spines are longer on the 
superomarginals and become progressively narrower along the borders of the 
inferomarginal plates. Each superomarginal plate carries a single pointed 
spine at its apex and 10 to 12 fasciole spinelets on each side. A single spine is 
present on most inferomarginal plates near to the upper edge. The terminal 
plates of this specimen are much abraded, but probably carry a single large spine 
either side of the terminal groove with a smaller spine set below and outside the 
former. 


a 
Fig. 2. Ctenodiscus sp. Mouth plate, proximal fascioles and adambulacral spinulation. 


The fascioles run across the actinal surface to the ambulacrum. The 
fasciole in the mid-line of the interradius divides behind the mouth plates 
(Fig. 2). The actinal plates between the first fascioles from the mid-line are 
made up of two rows of overlapping, sub-rounded plates. The other actinal 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wates, Vou. 96, Part 1 


52 SEA STAR GENUS CTENODISCUS 


plates are obscured by membrane. The adambulacral plates carry four to five 
furrow spines. Up to four spines may be present on the actinal surface of the 
adambulacral plates, amongst which a pair of spines may be distinct, especially 
near to the mouth. Occasional single spines are present on the actinal plates. 

The mouth plates are prominent, carry three or four oral spines and between 
10 and 12 suboral spines (Fig. 2). The innermost oral spines are the largest of 
that series. The second suboral spine is the largest of the suboral series. 


The tube feet are in two rows, pointed and without sucking discs. 


DISCUSSION 


The single specimen available is not adequate to confirm the existence of a 
further species of Ctenodiscus. However, some comparisons may be drawn with 
the species already ascribed to the genus. Sources of comparative information 
are given throughout the discussion. 

The paxillae of the specimen described are similar to those described as 
characteristic of C. procurator, being less than 0-5 mm. in height and without 
central spinelets (A. M. Clark, 1962). Fisher (1911) demonstrated the morpho- 
logical variability of C. crispatus and noted that the extremes of the paxillae and 
of body form of that species came within the variation shown by both C. procurator 
and C. australis. Fisher’s table of measurements showed that 16 marginal plates 
were present in C. crispatus at R of approximately 36 mm. The specimen under 
discussion has 16 marginal plates at about half that size. 


The number of furrow spines observed here comes well within the range given 
by Fisher for C. crispatus, i.e. three to five. 


C. orientalis is the only other form known with a large number of marginal 
plates (26-27 when R=52 mm.) and the number of fasciole spinelets are similar 
to the Tasmanian specimen. Comparison with Fisher’s (1919) figures of C. 
orientalis shows that the oral spinulation of the Tasmanian specimen differs 
from that species in that the second suboral spine is larger and there are more 
suboral spines (10-12 against 5-8). Fisher’s figures also show that central 
spinelets are present on the paxillae of C. ortentalis. 

On the basis of size and numbers of marginal plates, this specimen has been 
referred to Ctenodiscus orientalis Fisher. A similar Indo-Malay relationship 
has already been inferred by A. M. Clark (1962) for a species of Marginaster 
from deep water off eastern Tasmania. A number of hypotheses are available 
to fit the distribution observed. 

(1) All the attributed species of Ctenodiscus are but one species showing a 
wide range of morphological variation and bipolar distribution. 


(2) Two species of Ctenodiscus exist with north and south polar distributions. 


(3) The Tasmanian form shares its relationships with the tropical Indo-Malay 
fauna and is most closely related to Ctenodiscus orientalis. 


Without further material available, I have chosen the latter hypothesis. 
The specimen is also of interest as it adds another family to the known Australian 
asteroid fauna. 


References 


BERNASCONI, I., 1964.—Asteroideos argentinosos. Physis, 24 (68). 

D’yaxonovy, A. M., 1950.—Sea Stars (Asteroids) of the U.S.S.R. Seas, 1.P.8.T. edition. Jerusalem 
(1968). 

Cuark, A. M., 1962.—Asteroidea. B.A.N.Z. Antarctic Research Eaupedition Reports, B, ix. 

Fisuer, W. K., 1911.—Asteroidea of the North Pacific and adjacent waters. U.S. Nat. Mus. 
Bulletin, 76: 1. 

, 1919.—Starfishes of the Philippine Seas and adjacent waters. U.S. Nat. Mus. Bulletin, 

100: 3. 


PROCEEDINGS OF THE LINNEAN Socrety oF NEw SoutH WaAtzs, Vou. 96, Part 1 


THE GENUS NUNCIELLA ROEWER (OPILIONES, LANIATORES) 
WITH DESCRIPTION OF A NEW SPECIES FROM KANGAROO 
ISLAND, SOUTH AUSTRALIA 


Gao. ELuNT 
School of Zoology, University of New South Wales, Sydney 


[Accepted for publication 23rd September 1970] 


Synopsis 
The genus Nwnciella Roewer is redefined and the genus Cinuna Hickman is synonymized 
with it. A new species, N. kangarocensis n. sp., is described from Kangaroo Island, South 
Australia. A new key is given for the twelve species of Nunciella. The male genitalia of NV. aspera 
(Pocock), N. badia (Hickman), N. dentata (Hickman), N. tasmaniensis Hickman and N. tuberculata 
Forster are described. 


INTRODUCTION 


The genus Nunciella Roewer is one of the largest groups of triaenonychid 
harvestmen in Australia. Five species have been described from eastern. 
Australia south of Canberra, three from Tasmania, and two from the south-west 
of Western Australia. The new species from Kangaroo Island fills a gap in this 
distribution. 

The genus is of considerable interest because of the complex structure of the 
male genitalia, and sexual dimorphism in the chelicerae, pedipalps and coxae 
of leg 1. 

S Family TRIAENONYCHIDAE Soerensen, 1886 

Subfamily TRIAENONYCHINAE Pocock, 1903 
Tribe TRIAENONYCHINI Pocock, 1903 
Genus Nunciella Roewer, 1929 


Nunciella Roewer, 1929: 96; Roewer, 1931: 162; Forster, 1952: 26; 
Hickman, 1958: 35. 
Cinuna Hickman, 1958: 37, new syn. 

Although early authors have regarded the number of segments in tarsus 1 
a8 a reliable generic character, more recent work has shown that it must be used 
with caution in some genera (Goodnight and Goodnight, 1953). Forster (1954) 
allowed a variation of 2—4 tarsal segments in leg 1 for Nuncia and 5-7 for Hendea 
in his comprehensive revision of the New Zealand Laniatores. Recently, Forster 
(1965) described a cave species of Hendea with eight segments in tarsus 1, and 
remarked, ‘“‘ emphasis is placed on the tarsal formula for the separation of 
genera ...and it is of considerable interest to find that it may be influenced so 
readily by adaptation to a different environment”. Furthermore, Hickman 
(1958) has noted that variation (including asymmetry within an individual) 
ig common in some Australian genera. 

All the species of Nunciella described by Roewer possessed three tarsal 
segments in leg 1, a character he used in his description of the genus. Forster 
(1949), however, noted that the male of NV. tuberculata possessed four segments in 
tarsus 1, whilst the female had three. 

In his diagnosis of Nwnciella, Hickman (1958) retained Roewer’s diagnostic 
character of three segments in tarsus 1 while noting the situation in NV. tuberculata. 
On the basis of the segment number in tarsus 1 and the position of the eyemound, 
Hickman erected the closely related genus Cinuwna and placed in it C. badia 
(type species) and C. dentata from Tasmania. Oinuwna possessed four segments 


PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, Vou. 96, Part I 


54 GENUS NUNCIELLA ROEWER NEW SPECIES KANGAROO ISLAND 


in both sexes and an eyemound removed from the anterior margin of the carapace. 
The latter character is not valid because in males of NV. aspera and N. tuberculata 
the eyemound also rises from behind the anterior margin. 


It is obvious that the dimorphic variation in tarsus 1 in N. tuberculata 
(and WN. kangarooensis, n. sp.) causes problems in separating Cinuna from 
Nuneciella. Because of the similarity in the genitalia of the two genera (Figs. 
1-18) and the apparent absence of any clear-cut distinguishing characters, the 
limits of Nunciella must be extended to include species with four segments in 
tarsus 1. Thus Cinuna must be synonymized with Nuwneiella and the genus 
Nuneciella redescribed. 


Too little emphasis has been placed on the systematic importance of the 
male genitalia. Descriptions of genitalia have been given in only four of the 11 
previously described Nunciella species. Kauri (1955) described those in WV. aspera 
and WN. karriensis, and Forster (1955) those in N. montana and N. woolcock. 
Descriptions are given for species which have been examined by the author and 
whose genitalia had not been described previously. The penis of N. aspera 
is redescribed, as Kauri’s drawing is partly inaccurate. 


The genus Nunciella Roewer is now redefined as follows. 


Anterior margin of carapace unarmed or with row of tubercles or granules 
above. Eyemound low, generally rounded, rising from or behind anterior 
margin, unarmed except for small granules in some species. Tergites without 
spines or large tubercles, but with rows of granules. Chelicera of male with 
proximal retrodorsal boss on first segment. Pedipalp femur in male with a large 
modified proximoventral bifid tooth which is laterally flattened and directed 
backwards. With pedipalp lowered this tooth abuts another modified bifid 
tooth on coxa of leg 1. Calcaneus of each metatarsus much shorter than 
astragalus and that of leg 1 in male without notch below. Tarsal formula 3-4, 
8-15, 4, 4. Distitarsi 1 and 2 with two and mostly four segments respectively. 
Claw of both tarsus 3 and 4 three-pronged, middle prong larger than laterals. 
Apical portion of penis complex and usually compact. One superior and three 
inferior setae usual number on each ventral plate (N. tasmaniensis with two 
and five respectively). 


Type species: Nunciella aspera (Pocock). 


Nuneiella is closely related to the Tasmanian genus Nwncioides Hickman 
and the New Zealand genus Neonuncia Roewer. The males of these genera 
possess a strong proximal boss on the first segment of the chelicera, a highly 
modified bifid tooth placed proximoventrally on the pedipalp femur, and a 
calcaneus of leg 1 without notch below. Nunciella differs from Nwuncioides in 
having 3-4 segments rather than 5-6 in tarsus 1, and 4-5 compared with 6-8 
in distitarsus 2. In Neonuncia the scute, including the anterior margin, is smooth, 
without tubercles or granules, and there are only two inferior setae on each ventral 
plate of the penis. The structure of the genitalia and other characters suggest 
Nunciella is also related to Calliuncus Roewer. 

The 12 species in the genus Nwunciella may be distinguished by the following 
key. Females are not known for all species, so the key is based on male characters 
with the possible exception of NV. parvula Roewer, which Forster (1952) believes 
is a female, not a male as stated by Roewer (1931). NV. frontalis Roewer is 
omitted as it is probably a synonym of NW. aspera (Forster, 1952). 


Key to Species or NuNcIELLA (MALES) 


IeeTarsusnot lege with four sepmentsi> 4 a2 1aoote thie meee ne eased ei eh eee (2) 
Tarsusof ler swith) three segments:y. 4) hea 4 ae ie eee ae ee oe ae (5 

2. Distal portion of first segment of chelicera unarmed.................- N. tuberculata Forster 
Distal portion of cheliceral first segment with spine or prominent tubercle.......... (3) 

3. Pedipalp tibia with four strong retrolateral teeth........ NV. deniata (Hickman), new comb. 
Pedipalp tibia with three strong retrolateral teeth.................-...0 2.2 e eee 4 


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G. S. HUNT 55 


4. Pedipalp femur with five mediodorsal teeth...................... N. kangarooensis, n. sp. 
Pedipalp femur with four mediodorsal teeth.............. N. badia (Hickman), new comb. 

5. Prolateral surface of pedipalp femur with two distal teeth........................ (6) 
Prolateral surface of pedipalp femur without two distal teeth...................... (9) 

6. Prolateral surface of pedipalp patella armed sub-distally with strong tooth.......... (7) 
Prolateral surface of pedipalp patella armed sub-distally with small tubercle........ (8) 

7. Mediodorsal surface of pedipalp femur with four strong teeth.......... N. cheliplus Roewer 


Mediodorsal surface of pedipalp femur with four teeth, 1 and 3 very small, 2 and 4 strong.... 
Pee ye eMe reise toy siesiorte) ois cia) ohio (aliove: Ouetematey obs aeaye che su onencyt lays uckele, arsstelaterale N. granulata Roewer 
8. Penis with a rounded distally oriented swelling ventral to the inferior setae............ 
ee re aoe eee ale eee als ieuelinliesle, ovate Date. See ra cuicie, oye a elie ssley oie ela) Svenshualiey GeoRe N. karriensis Kauri 


Penis without a prominent swelling ventral to the inferior setae.......... N. aspera (Pocock) 
9. Femur of leg 3 with strong spine near proventral surface.............. N. woolcockt Forster 
Hemunsoimleresmwibhout suchiarspinek ieee er eee cee ae eicinicicie mele eiciciele eieie (10) 

10. Mediodorsal surface of pedipalp femur with five teeth................ N. parvula Roewer 
Mediodorsal surface of pedipalp femur with three teeth in proximal half.......... (11) 

11. Ventral surface of pedipalp femur with strong tooth in distal half and the prolateral surface 
RAIA ULTVAC Crate eee eke ara) e ahs) stay cus apes “sun cenalaia.cevsney eos austen ove ai save: sve eee N. tasmaniensis Hickman 
Ventral surface of pedipalp femur with tubercle in distal half and spinous tubercle at two- 
thirdssonyprolateralisurtaces....-- 5-142 ose cic ace le aid oi eile N. montana Forster 


Of the species in Nunciella, N. aspera and N. karriensis are from Western 
Australia, N. tasmaniensis, N. badia and N. dentata are from Tasmania, JN. 
cheliplus and N. parvula from Victoria, N. montana from New South Wales, 
N. woolcocki from the Australian Capital Territory, and N. kangarooensis, n. sp., 
from Kangaroo Island, South Australia. Roewer (1931) recorded N. granulata 
from New Zealand, but according to Forster (1954) it is probably an Australian 
species. 

Nuneciella aspera (Pocock) 
Triaenonyx aspera Pocock, 1903: 404; Loman, 1910: 133. 
Nuncia aspersa (Pocock): Roewer, 1915: 80; Roewer, 1923: 592. 
Nunciella aspera (Pocock): Roewer, 1929: 96; Forster, 1952: 27-29; Kauri, 
1955: 3-4. 
Nunciella frontalis Roewer, 1931: 163. 
Structure of Penis (Figs. 1-3): Penis complex and compact. Glans arising 
dorsally but curving ventrad, with terminal portion ventral (Fig. 2). Arising 
from glans, and flanking it on either side, two large curved plates. These plates 
present in all species, but relationship with glans not as evident in the eastern 
Australian forms. In these, the curved flanking plates meet dorsally and 
presumably obscure the line of fusion with the glans. The glans and flanking 
plates constitute the aedeagus. 

Ventral plates fused and enlarged basally, but not with a pronounced 
distally-oriented swelling ventral to inferior setae. Plates tapering rapidly 
distal to inferior setae, separated by deep notch. Hach ventral plate with one 
superior and three inferior setae. 

Material Examined: Ten males, three females (type material not examined). 


Locality: Strong’s Cave, near Margaret River, Western Australia: under loose 
rocks and debris in entrance doline. Coll. G. S. Hunt 5.1.65. 


Nunciella badia (Hickman), new comb. 

Cinuna badia Hickman, 1958: 38-40. 

Structure of Penis (Figs. 4-6): Apical portion complex and compact. Glans 
rugose and swollen sub-apically (Fig. 4), protected dorsally by curved hood-like 
process. Two curved plates dorsal to hood, fused along mid-dorsal line except 
for narrow cleft. Each ventral plate with one superior and three minute inferior 
setae. Distal edge of plate serrated and distodorsal region rugose (Fig. 5). 
Plates fused except for shallow indentation. 


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56 GENUS NUNCIELLA ROEWER NEW SPECIES KANGAROO ISLAND 


—— 
O.1 MM 


Text-fig. 1 
Figs. 1-3. Nunciella aspera (Pocock). 1, Apical portion of penis, ventral view ; 2, lateral 
view; 3, dorsal view. 


Figs. 4-6. Nunciella badia (Hickman), new comb. 4, Apical portion of penis, ventral view ; 
5, lateral view; 6, extreme apical portion, dorsal view. 


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G. 8S. HUNT 57 


IO 


Text-fig. 2 
Figs. 7-9. Nunciella dentata (Hickman), new comb. 7, Apical portion of penis, dorsal 
view; 8, lateral view; 9, ventral view. 


Figs. 10-12. Nuwnciella tasmaniensis Hickman. 10, Apical portion of penis, dorsal view ; 
11, lateral view; 12, ventral view. 


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58 GENUS NUNCIELLA ROEWER NEW SPECIES KANGAROO ISLAND 


Material Examined: Holotype male and allotype female. 

Localities : Holotype male from near Lake St. Clair, Tasmania: under rotting 
log. Coll. V. V. Hickman 7.11.45. Australian Museum, Sydney, Cat. No. 
K68133. Allotype from side of Lyell Highway, 23 miles from Queenstown, 
Tasmania: under rotting log. Coll. V. V. Hickman 23.v.54. Australian 
Museum, Sydney, Cat. No. K68134. 


Nunciella dentata (Hickman), new comb. 
Cinuna dentata Hickman, 1958: 40-42. 
Structure of Penis (Figs. 7-9): Apical portion complex and compact. Glans 
apparently lying between two curved plates, appearing narrow and tubular with 
an apical expansion. Curved plates flared out laterally, fused along mid-dorsal 
line except for indentation. Ventral plates fused except for wide smoothly 
curved shallow indentation, each with one superior and three inferior setae. 


Material Examined: Holotype male. 
Locality: Slopes of Mt. Wellington, Tasmania. Coll. C. Oke Sept., 1949. 
Australian Museum, Sydney, Cat. No. K68135. 


Nunciella tasmaniensis Hickman 
N. tasmaniensis Hickman, 1958: 35-38. 
Structure of Penis (Figs. 10-12): Apical portion complex. Glans laterally 
flattened, narrowed sub-apically, directed slightly ventrally and striated sub- 
distally with lateral ridges (Fig. 11). Flanking plates wing-like, joining medio- 
ventrally and mediodorsally. Ventral plates fused except for wide sharply 
cornered indentation, each with two strong superior and five strong inferior setae. 


Material Examined: Holotype male and allotype female. 


Locality: Weldborough Pass, Tasmania: under rotting logs. Coll. V. V. 
Hickman 25.viii.53. Australian Museum, Sydney, Cat. No. K68132. 


Nunciella tuberculata Forster 
N. tuberculata Forster, 1949: 71, 73-75, 78. 
Structure of Penis (Figs. 13-15): Glans ventrally placed and narrowed apically. 
Flanking plates large, flared out laterally and fused along mid-dorsal line except 
for very deep and narrow notch. Ventral plates fused except for a sharply 
cornered indentation, each with one superior and three inferior setae. Portion 
of plates distal to inferior setae relatively enlarged. 
Material Examined: Holotype male, allotype female and four paratype males. 
Localities: Holotype, allotype and three paratypes from Diamond Creek, 
Victoria. Coll. J. A. Kershaw 11.iv.05. Paratype male from Kallista, Victoria : 
under log. Coll. A. N. Burns 18.ix.46. National Museum of Victoria, Melbourne. 


Nunciella kangarooensis, ni. sp. 
Holotype male. 
Measurements (mm.): Length scute 3-90, width scute 3-62, length carapace 
1-64, length genital operculum 0-43, width genital operculum 0-42. 


Cox. Troch. Fem. Pat. Tib. Met. Tar. Total 
Leg 1 BS, 1-44 0-68 1-96 0-91 1-52 1-95 1-28 9-74 
Leg 2 we 1-99 0-74 2-74 1-15 2-31 2°77 _ 2°70 14-40 
Leg 3 ss 1-49 0-78 2-02 0-85 1-54 2-13 1°35 10-16 
Leg 4 & 1-96 0-94 2-77 1-18 2-16 3°34 1-60 13-95 


Pedipalp Jn oils 0-61 2-02 1-16 1-42 — 1-42 7-54 
Chelicera : First segment 1-55, second segment 1:59, total 2-88. 


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G. S. HUNT 


> 


west ort més en 


l6 7 


— 
O.1 MM 
Text-fig. 3 


Figs. 13-15. Nunciella tuberculata Forster. 13, Apical portion of penis, ventral view : 


14, lateral view; 15, dorsal view. 
Figs. 16-18. Nuncvella kangarooensis, n. sp. 16, Apical portion of penis, dorsal view : 
17, lateral view; 18, extreme apical portion, ventral view. 


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60 GENUS NUNCIELLA ROEWER NEW SPECIES KANGAROO ISLAND 


Colour: Dorsal body pattern as in Fig. 19. Carapace with background colour 
orange-brown superimposed with darker red-brown markings, including 
reticulations lateral to eyemound and distinct markings posterior to eyemound. 
Tergites generally darker than carapace, overall colour dark brown to black. 
Lighter markings down sides. Prominent club-shaped stripe posterior to scutal 
groove. Yellowish circular patch surrounding each of the two mesial granules 
in areas 2 and 3, less distinct patches in areas 1, 4 and 5. Row of patches along 
each free tergite. Pedipalps and chelicerae orange-brown with darker reticula- 
tions. Legs yellow-brown with dark brown and black markings. 


Body (Figs. 19, 21-23): Eyemound low, less than half as high as long, directed 
slightly forward, rising abruptly from just behind anterior margin of carapace 
and sloping back gently posterior to apex (Figs. 21-22). Apex with small setose 
granule. Six small tubercles directed obliquely forward and some small granules 
along anterior margin of carapace. Five other processes lower on anterior 
margin: one mesially placed and curved slightly upwards, one abutting retro- 
proximal boss of each chelicera, one outside each pedipalp. Each tergal area 
and free tergite with transverse row of setose granules, mesial ones larger. Most 
of scute with minute granulations but free tergites more sparsely covered. Scutal 
groove prominent mesially, groove between areas 4 and 5 distinct, extending 
towards lateral margins, that between areas 1 and 2 faintly indicated mesially, 
others indistinct. Free tergites tending to be fused to each other and to scute, 
a condition approaching that of NV. tasmaniensis. 


Sternites each with row of tiny setose granules. Margin of spiracle only 
obscured by one or two tubercles on coxa 4. Genital operculum smooth except 
for scattered hairs. 


Pems (Figs. 16-18): Apical portion complex and compact. Glans ventrally 
placed and dorsoventrally flattened, with pair of curved plates dorsal to it. 
Plates fused mid-dorsally except for large V-shaped notch. Ventral plates 
fused except for broad shallow indentation, each with one superior and three 
inferior setae. 


Chelicerae (Figs. 19, 27): Strong retroproximal boss on dorsal surface of first 
segment abutting process on anterior margin of carapace. Dorsal surface with 
a few small granules and prodistal spine directed obliquely forward. Dorsal 
surface of second segment with small inner sub-proximal spine, retrodorsal 
tubercle at two-thirds and small spine at retrodistal corner. 


Pedipalps (Figs. 24, 25): Moderately large. Trochanter with a few granules: 
Dorsal surface of femur with row of five prominent teeth, third and fourth largest: 
On ventral surface median proximal tooth modified into large laterally compressed 
process directed obliquely backwards (Fig. 25). With pedipalp lowered, posterior 
edge of this process in contact with modified cup-like process on coxa l. Four 
other teeth along ventral surface, second and fourth strong, first and third weak. 
Two large sub-distal teeth low on prolateral surface. Granule at two-thirds 
high on prolateral surface enlarged into denticle. Patella with large sub-distal 
tooth low on prolateral surface. Tibia with three teeth down each ventrolateral 
margin. Tarsus with small denticle followed by three teeth down retroventral 
margin and three teeth down proventral margin. Claw about 0-25 length 
of tarsus. 


Legs: Coxa 1 with large cup-like process in prodistal corner (Figs. 23, 29) and 
two spines near proventral margin, more distal stronger. Retroventral edge 
with sub-distal tubercles. Coxa 2 with small sub-distal spine on retrolateral 
surface and small tubercles more proximally. Maxillary lobe with three setose 
tubercles. Coxa 4 with pro- and retrolateral tubercles. Trochanter of each leg 
with small tubercles or granules, femora with rows of small tubercles and/or 
granules, those on leg 1 ventral surface largest. Each calcaneus much shorter 


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O.5 MM 


1.O MM 


Text-fig. 4 


Figs. 19-23. Nunciella kangarooensis, n. sp. 19, Dorsum of male showing colour pattern, 
with first segment of chelicerae also shown; 20, dorsum of female showing colour pattern, with 
first segment of chelicerae also shown; 21, male body, lateral view; 22, eyemound of male, 
lateral view ; 23, sterno-coxal region in male. 


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62 SENUS NUNCIELLA ROEWER NEW SPECIES KANGAROO ISLAND 


oe! 
O.2MM 


Text-fig. 5 
Figs. 24-29. Nunciella kangarooensis, n. sp. 24, Male pedipalp, prolateral view > 
25, retrolateral view; 26, female pedipalp, retrolateral view; 27, second segment of male 
chelicera, dorsal view ; 28, process in prodistal corner of coxa 1 in female; 29, same process 
in male. 


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G. S. HUNT 63 


than astragalus, calcaneus of leg 1 without notch. Distitarsus of leg 1 two- 
segmented, that of leg 2 four-segmented. Tarsal formula: 4, 9-13, 4, 4. 
Allotype female. 


Measurements (mm.): Length scute 3-89, width scute 3-45, length carapace 
1-49, length genital operculum 0-51, width genital operculum 0-47. 


Cox. Troch. Fem. Pat. Tib. Met. Tar. Total 
Leg 1 ae 1-18 0:57 1-82 0-81 1-35 1:78 1-08 8-59 
Leg 2 ae 1:66 0:74 2-44 0-98 2-08 2:53 2-43 12-86 
Leg 3 os Io B37 0:59 1:76 0-78 1-35 1-99 1322 9-06 
Leg 4 ay 1-72 0-91 2-52 1:02 1:99 3°04 ets y55 12-75 
Pedipalp sa 1:01 0:54 1-62 0-88 1-15 — 1-08 6-28 

Chelicera: First segment 1-04, second segment 1-66, total 2-7C. 
1.8 


CARAPACE LENGTH (MM) 


0.80 0.85 0.90 O95 1.00 


RATIO SCUTE WIDTH TO LENGTH 


Text-fig. 6 


Fig. 30. Nunciella kangarooensis, n. sp. Range of stoutness (ratio scute width to length) 
and size segregation of large and small males (dots) and females (circles). 


Female Characteristics: As in other species in the genus, sexual dimorphism is 
strong. Female differs from male in the following respects. Dorsal pattern 
without median stripe, tergal areas more clearly defined, yellow patches around 
granules more prominent (Fig. 20). Retroproximal boss on dorsal surface of 
cheliceral first segment absent, but small tubercle occupies a near position ; 
corresponding process on anterior margin reduced. Spine in retrodistal corner 


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64 GENUS NUNCIELLA ROEWER NEW SPECIES KANGAROO ISLAND 


of second segment absent. Pedipalps (Fig. 26) smaller. Proximoventral tooth 
on femur not extremely modified, coxa 1 without extreme cup-like modification 
to bifid process in prodistal corner (Fig. 28). Body less stout than male, though 
not in all the population (Fig. 30). Female type of similar length to male, though 
females in general shorter (Fig. 30). Degree of fusion of free tergites less. As 
with NV. tuberculata, tarsus 1 in the female with three segments. Tarsal formula : 
3, 8-11, 4, 4. 
REMARKS 


The males can be separated from those of other species in the genus, except 
some individuals of NV. aspera and N. karriensis, by the presence of five medio- 
dorsal teeth and two strong prolateral teeth on the pedipalp femur. A less 
eranulate eyemound distinguishes them from the other two species. 


N. kangarooensis, n. sp. is similar to N. twberculata in having four segments 
in tarsus 1 of the male and three in the female. It resembles many other species 
in having a club-shaped median stripe on the dorsal surface of many males. It 
resembles NV. tasmaniensis in the tendency for fusion of the free tergites in males. 
The structure of the genitalia suggests N. kangarooensis is more closely related 
to the eastern Australian species than those in Western Australia. 


Types 

Holotype male, allotype female and two paratype males in the Australian 
Museum, Sydney, Cat. No.’s K68773, K68774 and K68775. One male and 
one female paratype in the South Australian Museum, Adelaide. 


Locality 

Near Shackle Road crossing of Rocky River, Flinders Chase, Kangaroo 
Island, South Australia: in moist litter amongst the dead drooping blade-like 
leaves of tall monocots, one to two metres high (Type Locality). The habitat 
is Shaded by thick vegetation beside the river. Altitude, under 60 metres. Coll. 
G. 8S. Hunt 7.1.69. 


Variations 

The collection contains 25 males, 23 females and one small nymph. As can 
be seen from Fig. 30, the males fall into two distinct groups based on size, for 
which carapace length is used as an index. The females show less tendency to 
fall into groups, the majority lying between the two groups of males in size. The 
Size variations are summarized in Table 1. 


TABLE | 
Size Variation in Nunciella kangarooensis, n. sp. 


Scute length Seute width Carapace length 
Number (mm.) (mm.) (mm.) 
of 
Specimens Range Mean Range Mean Range Mean 
Group of larg 
males .. a6 17 3:69-4:42 3-93 3:24-3:82 3-56 1:50-1:72 1-67 
Group of small 
males .. si 8 2:86-3:21 3-03 2-40-3-11 2-73 1-18-1-35 1-29 
Females me 21 3:01-3:89 3-53 2-77-3-45 3-18 1-32-1-57 1-48 


Two small females are not treated in the table as they do not fall into the 
main group. They have a scute length of 2-70 and 2:85 mm., scute width of 
2-64 and 2:30 mm., and carapace length of 1-11 and 1-22 mm. The measure- 
ments of the nymph are 2-53, 2-09 and 1-04 mm. respectively. 

Both males and females show roughly the same range in stoutness (ratio 
scute width to length, Fig. 30). Individuals vary in the tendency for fusion of 


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G. 8S. HUNT 65 


the free tergites, the group of small males and the females generally showing less 
tendency. Both sexes vary slightly in body pattern. The density of granules 
on the scute varies, and the strength of teeth and shape of eyemound is subject 
to minor variation. The number of segments in tarsus 1, 3 and 4 is constant 
for each sex, except one male with five, not four, in right tarsus 1. The number 
of segments in tarsus 2 is subject to asymmetry and wide variation. The mean 
in females is 9-6 compared with 10-5 in males. The mode, however, is 10 in 
both sexes. The number of segments in distitarsus 2 is four in both sexes, except 
for one male with five in the right distitarsus. 


Male Dimorphism 

The two distinct groups of males (Fig. 30) are of interest. Individuals in 
both groups seem to have secondary sexual characteristics developed to the 
same degree. The two groups may indicate male dimorphism which has been 
recorded by Forster (1954) in some New Zealand triaenonychids and which 
also occurs in some Australian genera, notably Hquitius Simon. The presence 
of two small females in the collection means that no definite conclusions can be 
made until the life history of the species is known and more specimens studied. 


Habitat 

Like most of the Laniatores, NV. kangarooensis, n. sp. lives in humid situations 
which are provided by the shaded areas beside Rocky River. The species 
probably occurs beside other streams and around the edges of lagoons on the 
island. No specimens were found during limited searching on higher ground 
under the dead drooping leaves of the Xanthorrhoea (yakka) grass-tree, or under 
rocks in dolines on the limestone. Both situations, however, are potentially 
favourable habitats for harvestmen. 


ACKNOWLEDGEMENTS 


I wish to thank members of the Australian Speleological Federation, whose 
expedition to Kangaroo Island provided an opportunity for specimen collection. 
I am indebted to Mr. J. McNally, Director of the National Museum of Victoria, 
for the loan of material, and to Dr. C. N. Smithers and Mr. M. R. B. Gray of 
the Australian Museum for the assistance they have given me. I am especially 
grateful to Dr. A. M. Richards, University of New South Wales, and Dr. R. R. 
Forster, Director of the Otago Museum, New Zealand, for reading through the 
manuscript and offering useful criticisms and advice. 


_References 


Forster, R. R., 1949.—Australian Opiliones. Mem. Nat. Mus. Vict., No. 16: 59-89. 
, 1952.—Western Australian Opiliones. JI R. Soc. West. Aust., 36: 23-29. 
, 1954.—The New Zealand harvestmen (sub-order Laniatores). Canterbury Mus. Bull., 
No. 2: 1-329. 
, 1955.—Further Australian Opiliones. Aust. J. Zool., 3 (3): 354-411. 
, 1965.—Harvestmen of the suborder Laniatores from New Zealand caves. Rec. Otago 
Mus., No. 2: 1-18. 
Goopniecut, C. J., and Goopnicut, M. L., 1953.—Taxonomic recognition of variation in Opiliones. 
Syst. Zool., 2 (4): 173-180. 
Hickman, V. V., 1958.—Some Tasmanian harvestmen of the family Triaenonychidae. Pap. 
Proc. R. Soc. Tasm., 91: 1-116. 
Kauri, H., 1955.—Report from Professor T. Gislen’s expedition to Australia i 1951-1952. 
9. Harvest-spiders from §8.W. Australia. Lunds. Uni. Arsskr. N.F., Avd. 2, 50 (11): 1-10. 
Loman, J. C. C., 1910.—Opiliones. Die Fauna Stidwest-Australiens, 3 (4): 127-134. 
Pocock, R. I., 1903.—On some new harvest-spiders of the order Opiliones from southern continents. 
Proc. Zool. Soc. Lond., 1902, Part 2: 392-413. 
Rorewer, C. Fr., 1915.—Die Familie der Triaenonychidae der Opiliones Laniatores. Arch. 
Naturg., Berlin, 1914, 80a (12): 61-168. 
, 1923.—Die Weberknechte der Hrde. G. Fischer, Jena, 1116 pp. 
, 1929.—Contribution to the fauna of Rottnest Island. No. V. Opiliones in the 
Western Australian Museum. J/ R. Soc. West. Aust., 15: 95-98. 
, 1931.—Uber Triaenonychidae (VI Erganzung der ‘‘ Weberknechte der Erde ”’, 1923). 
Zetts. f. wiss. Zool., 138 (1): 187-185. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SourH Wates, Vout. 96, Part I 


LATE DEVONIAN CONODONTS FROM THE LUTON FORMATION, 
NORTHERN NEW SOUTH WALES 


G. M. Puimip AND J. H. JACKSON 
Geology Department, University of New England, Armidale, N.S.W. 


(Plates v, v1) 


[Accepted for publication 18th November 1970] 


Synopsis 
A small Upper Devonian conodont fauna is described from a limestone lens in the lower part 
of the Luton Formation, northern New South Wales. The fauna indicates a late Fammenian 
(to VI) age. 


INTRODUCTION 


In this article a small conodont fauna of late Fammenian age from the base 
of the Luton Formation, northern New South Wales, is described. The occurrence 
of conodonts in an impure limestone horizon in the Late Devonian rocks of this 
region promises to provide an accurate correlation of these strata, as well as 
locating more precisely the Devonian/Carboniferous boundary in northern 
New South Wales. To date correlation of the thick Late Devonian epiclastic 
sequences of the ‘‘ Tamworth Trough” has been based on the few isolated 
occurrences of poorly preserved ammonoids (Pickett, 1960; Jenkins, 1966, 
1968). 

This work is an aspect of a project on eastern Australian Devonian bio- 
stratigraphy, supported by the Australian Research Grants Committee. We 
are grateful to Dr. Gilbert Klapper, University of Iowa, for useful discussions 
of this paper. 


LOCALITY AND SIGNIFICANCE OF FAUNA 


The conodonts were recovered from an impure limestone lens in 
the lower part of the Luton Formation, 74 miles south-south-west of Horton 
(150° 24-3’ B., 30° 14-8’S.). This falls on the Horton Map Sheet provided by 
McKelvey and White (1964), who also gave a summary of the stratigraphy. 
These writers provisionally placed the base of the Carboniferous at the base of 
the Luton Formation. To date no other horizons in the Luton Formation have 
yielded conodonts. 

In all, approximately 50 kg. of sandy limestone was processed, and this 
yielded some 250 identifiable conodonts. The conodonts are dark brown in 
colour and have retained their ‘‘ white matter’’. Most of the conodonts are 
broken. The fauna is dominated by platform elements, in particular Palmatolepis 
gracilis, Polygnathus vogest, Polygnathus cf. pennatulus and Pseudopolygnathus 
dentilineatus. Twenty-six different species of disjunct conodont were recognized. 

Ziegler (1962) has provided the standard conodont zonation of the Upper 
Devonian, on the basis of sections mainly in the Rhenish Schiefergebirge. This 
was related by Ziegler to the well-established German ammonoid succession, 
and has been found to apply not only in Europe, but elsewhere in the world 
[e.g.. in N. America (Klapper, 1966) and W. Australia (Glenister and Klapper, 
1966) ]. 


The highest Fammenian zone was named the costatus Zone by Ziegler (1962) 
and divided into three on the basis of the ranges of subspecies of Spathognathus 
costatus sensu Ziegler and Palmatolepis gonioclymeniae. The Lower costatus 


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G. M. PHILIP AND J. H. JACKSON 67 


Zone falls in the upper part of the Clymenia Stufe (to V), whereas the Middle 
and Upper costatus Zones are approximately coextensive with the Wocklumeria 
Stufe (to VI). 

Although it is clear that the Luton conodont fauna is that of the costatus 
Zone, in the absence of S. costatus sensu Ziegler and P. gonioclymeniae it is difficult 
to arrive at a more precise estimate of its position. 


Palmatolepis gracilis sigmoidalis ranges throughout the costatus Zone. 
Polygnathus vogesi is said by Ziegler (1962) to range from the Middle costatus 
Zone into the Gatiendorfia Stufe. Pseudopolygnathus trigonicus was given by 
Ziegler (1962) as occurring in the Middle and Upper costatus Zones, although 
Freyer and Zakowa (1967) have extended the range down into the Lower costatus 
Zone. Spathognathodus aculeatus sensu Ziegler is usually considered to occur 
in the Lower and Middle costatus Zones, but there are problems in the identi- 
fication of this form species. Another platformed element in the fauna suggestive 
of a to VI age is Polygnathus sp. cf. symmetricus. Other elements appear to be 
longer ranging. 

The evidence therefore favours reference of the Luton conodont fauna to 
the Middle costatus Zone, equivalent to the early Wocklumeria Stufe (to V1). 
This indicates that the Devonian-Carboniferous boundary in New South Wales 
is to be found in the upper part of the Luton Formation. The to VI age for the 
lower part of the Luton Formation lends strong support to Pedder’s (1968) 
conclusion that the Kiah Limestone Member (with Cymaclymenia borahensis 
Pickett) of the underlying Mandowa Mudstone is of te V age. 


SYSTEMATICS 
Genus BRYANTODUS Bassler, 1925 


Type species: Bryantodus typicus Bassler, 1925. 


Bryantodus sp. (Pl. v, Fig. 1) 

Figured hypotype: ¥F10901/1. 
Remarks: The most complete fragment of this form lacks the posterior bar. 
The anterior bar is arched and gently flexed, with fused, laterally compressed 
denticles. As far as can be ascertained, the species represented is similar to 
Bryantodus typicus Bassler (Huddle, 1968, p. 11, Pl. 3, figs. 1-15; PI. 4, figs. 
12-15). 

Genus DIPLOPODELLA Bassler, 1925 
Type species: Diplopodella bilateralis Bassler, 1925. 


Diplopodella sp. (Pl. v, Fig. 16) 
Figured hypotype: ¥10901/16. 


Remarks: The more complete of the two available specimens lacks one lateral 
bar. The cusp is laterally compressed and projects posteriorly to form a short 
posterior bar bearing two compressed denticles. The lateral limbs are deep and 
bear closed spaced, fused denticles. There is no basal cavity. 


Genus HIBBARDELLA Bassler, 1927 
Type species: Prioniodus angulatus Hinde, 1879. 


Remarks: Recent discussions of the application of the genera Aibbardella, 
Roundya Hass, Ellisonia Miller and Diplopodella Bassler have been given by 
Philip (1966), Miiller and Clark (1967) and Huddle (1968), who are all in substantial 
agreement. Here Huddle’s interpretation of these genera has been followed. 
More recently Rhodes ef al. (1969, p. 111) have proposed a new subgenus 
Hassognathus, based on Trichognathus separatus Branson and Mehl, for similar 
forms in which the basal cavity is located more beneath the posterior bar. They 
also maintain Roundya as a subgenus of Hibbardella. 


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68 LATE DEVONIAN CONODONTS, LUTON FORMATION, N.S.W. 


Hibbardella sp. (Pl. v, Fig. 3) 
Figured hypotype: F10901/3. 
Remarks: The single available specimen of this form has a large subquadrate 
cusp and a deep basal cavity. It closely resembles Hibbardella barnetiana 
(Hass, 1953, p. 89, Pl. 16, figs. 8-9), the type species of Roundya Hass, but as all 
three processes are broken, no positive identification is made. 


Genus HINDEODELLA Bassler, 1925 
Type species: Hindeodella subtilis Bassler, 1925. 


Hindeodella sp. (Pl. v, Fig. 2) 

Figured hypotype: F10901/2. 

Remarks: Specimens of Hindeodella in the collection are all extremely frag- 
mentary. It is likely that the specimen illustrated in Pl. v, Fig. 2 is referrable 
to H. subtilis Bassler as interpreted by Huddle (1968, pp. 17-18). The specimen 
illustrated as Hindeodella ? sp. in Pl. v, Fig. 11 (F10901/11) is apparently a 
different species, and, in the strong downward deflection of the anterior bar, 
resembles more closely Hindeodella alternata Ulrich and Bassler (Huddle, 1968, 
pp. 15-16, Pl. 5, figs. 1,3). As the posterior bar is missing, however, the specimen 
cannot be positively assigned to Hindeodella. 


Genus LIGONODINA Bassler, 1925 
Type species: Ingonodina pectinata Ulrich and Bassler, 1926. 


Tigonodina bicurvata Mound (Pl. v, Fig. 10) 
LTigonodina delicata Branson and Mehl, 1934, p. 199, Pl. 14, figs. 22, 23. 
Ligonodina bicurvata Mound, 1968, p. 489, Pl. 66, figs. 22, 30 (q.v. for full 
synonymy). 
Tigonodina beata Rhodes, Austin and Druce, 1959, p. 135, Pl. 26, figs. 4-6). 
Not Phragmodus delicatula Branson and Mehl —Ligonodina delicata (Branson and 
Mehl). 
Figured hypotype: F10901/10. 
Description: This is a delicate species of Ligonodina with a long, gently recurved, 
somewhat flattened cusp continued downwards and inwards as a long anticusp. 
Up to about five isolated rounded denticles on anticusp; posterior bar with 
slender, backwardly directed, well separated, rounded denticles. Aboral surface 
of posterior bar marked with a longitudinal groove, which opens into the large 
basal cavity and is continued to the extremity of the anticusp. 
Remarks: Mound’s (loc. cit.) correction of the primary homonym which arose 
from Bergstrém’s (1964, pp. 28-29) reference of Phragmodus delicatus Branson 
and Mehl to Ligonodina has priority over that of Rhodes et al. Ethington (1965) 
has noted that the L. bicurvata ranges from the Middle Devonian through into 
the Early Carboniferous. 


Genus LONCHODINA Bassler, 1925 
Type species: Lonchodina typicalis Bassler, 1925. 


Lonchodina spp. (Pl. v, Figs. 9, 14-15) 
Figured hypotypes : F10901/9, 14, 15. 
Remarks: A number of asymmetrical, twisted and arched bar elements are 
present in the collection. Three different species of Lonchodina appear to be 
represented by the illustrated specimens. Of these, that shown in PI. v, Fig. 15 
resembles closely Lonchodina clavata (Hinde). However, more material is 
necessary before positive identification can be made. 


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G. M. PHILIP AND J. H. JACKSON 69 


Genus NEOPRIONIODUS Rhodes and Miller, 1956 
Type species: Prioniodus conjunctus Gunnell, 1931. 


Neoprioniodus oligus (Cooper) (Pl. vi, Fig. 14) 
Prioniodus oligus Cooper, 1939, p. 405, Pl. 46, figs. 9-11, 63, 71 ; Pl. 47, figs. 20-21. 
Prioniodina oliga (Cooper), Bischoff, 1957, pp. 47-48. 
Neoprioniodus oligus (Cooper), Hass, 1959, p. 384, Pl. 47, fig. 3. 
Figured hypotype: F10900/14. 
Description: A form of Neoprioniodus in which the denticles of the posterior 
bar are strongly compressed and may be completely fused to form a deep ridge 
which merges with the cusp. Cusp flattened with anterior and posterior keels. 
Basal cavity large with a flaring inner margin. 
Remarks: The single specimen recovered from the Luton Formation is identical 
with the material described by Hass (loc. cit.) from the Chappel Limestone of 
Texas. Hitherto the species appears to have been recorded only from Early 
Carboniferous strata. 


Genus OZARKODINA Branson and Mehl, 1933 
Type species: Ozarkodina typica Branson and Mehi. 


Ozarkodina spp. (Pl. v, Figs. 4, 12, 13) 

Figured hypotypes: F10901/4, 12, 13. 

Remarks: Three separate species of Ozarkodina appear to be present in the 
collection, but all of the specimens are fragmentary. The form illustrated in 
Pl. v, Fig. 4 resembles closely O. macra Branson and Mehl, whereas that shown in 
Pl. v, Fig. 12 is a more delicate unit, and so may be compared with O. elegans 
(Stauffer). The specimen illustrated in Pl. v, Fig. 13 has a tiny basal cavity 
and a poorly differentiated cusp. These characters appear to distinguish it 
from previously described species. 


Genus PALMATOLEPIS Ulrich and Bassler, 1926 
Type species: Palmatolepis perlobata Ulrich and Bassler, 1926. 
Remarks: Attempts to subdivide this widespread Devonian form genus (Miiller, 
1956 ; Helms, 1963) have not met with general acceptance. Although the genus 
is generally considered to be confined to the Late Devonian, Huddle (1968, p. 33) 
has suggested that his (1934) P. minuta (non Branson and Mehl) is an authentic 
Early Mississippian species. 


Palmatolepis graciiis Branson and Mehl 
Palmatolepis gracilis Branson and Mehl, 1934, p. 238, Pl. 18, figs. 2, 8 (not fig. 5) ; 
Mehl and Ziegler, 1963, pp. 200-205, PI. 1, figs. 1-2. 
Palmatolepis (Deflectolepis) deflectens Muller, 1956a, p. 32, Pl. 11, figs. 28-39. 
Remarks: This species of Palmatolepis usually possesses a narrow platform 
running parallel with the carina. The platform is widest anteriorly and tapers 
to the posterior end. The outer margin of platform is regularly convex at the 
anterior end, and the inner margin forms an anterior subcircular lobe. Ziegler 
(1962) distinguished two subspecies, both of which occur in the Luton Formation. 
Palmatolepis gracilis gracilis Branson and Mehl (Pl. v1, Figs. 8-10, 26-29) 
Palmatolepis gracilis Branson and Mehl, 1934, p. 238, Pl. 18, figs. 2, 8 (not fig. 5) ; 
Wolska, 1967, p. 395, Pl. 11, figs. 1-3, 5 (not fig. 4 =P. gracilis sigmoidalis). 
Palmatolepis deflectens deflectens Muller, Ziegler, 1962, p. 56, Pl. 3, figs. 17-22 ; 
Freyer in Freyer and Zakowa, 1967, pp. 114-115, Pl. 1, fig. 1; Pl. 2, figs. 3-4. 
Palmatolepis gracilis gracilis Branson and Mehl, Klapper, 1966, p. 31, Pl. 6, fig. 3 ; 
Klapper and Glenister, 1966, pp. 814-815, Pl. 90, fig. 6 (q.v. for full 
synonymy); Druce, 1969, pp. 88-89, Pl. 17, figs. 3a—5d. 


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70 LATE DEVONIAN CONODONTS, LUTON FORMATION, N.S.W. 


Figured hypotypes : F10900/8-10, 26-29. 

Description: This subspecies of P. gracilis has a gently to strongly curved blade 
and carina. The keel is offset parallel to the margin of the inner platform lobe 
to form a semicircular ridge (Pl. vi, Fig. 27). 

Remarks: Klapper (loc. cit.) has pointed out that in some specimens of P. gracilis 
gracilis the platform may become relatively large so that they resemble P. gonio- 
clymeniae. In this latter species, however, the posterior end is arched upwards, 
the carina is more sharply deflected, and this deflection takes place closer to the 
anterior end. The larger specimens in the Luton fauna have been previously 
listed as P. gonioclymeniae in Pedder (1968, p. 141). 


Palmatolepis gracilis sigmoidalis Ziegler (Pl. vi, Figs. 5-7) 
Palmatolepis deflectens sigmoidalis Ziegler, 1962, pp. 56-57, Pl. 3, figs. 24-28. 
Palmatolepis gracilis sigmoidalis Ziegler, Klapper, 1966, p. 31, Pl. 6, fig. 8 (q.v. 

for full synonymy) ; Druce, 1969, p. 89, Pl. 17, figs. 2a-b. 
Palmatolepis gracilis Branson and Mehl, Wolska, 1967, p. 395, Pl. 11, fig. 4 (not 
figs. 1-3, 5 =P. gracilis gracilis). 
Figured hypotypes: ¥10900/5-7. 
Description: In this subspecies of P. gracilis the carina and platform are strongly 
sigmoidal. 
Remarks: The specimens from the Luton Formation bear out Klapper’s (loc. 
cit.) observation that this subspecies lacks the semicircular offset of the keel 
which is present in the nominate subspecies. P. gracilis sigmoidalis is confined to 
the Spathognathodus costatus Zone (sensu Ziegler, 1962) in both Europe and 
N. America. 


Genus POLYGNATHUS Hinde, 1879 
Type species: Polygnathus dubius Hinde, 1879. 


Polygnathus cf. pennatulus Ulrich and Bassler (Pl. v, Figs. 23-27) 

Cf. Polygnathus pennatulus Ulrich and Bassler, 1926, p. 45, Pl. 7, fig. 8, text-fig. 
i () 

Polygnathus pennatula Ulrich and Bassler, Huddle, 1968, pp. 40-41, Pl. 15, figs. 
11-12 (q.v. for full synonymy). 

Figured hypotypes: ¥F10901/23-27. 


Description: The platform is thick and subtriangular with a high carina flanked 
by short, deep troughs. The unit is somewhat bowed and arched. The margins 
of the platform are crenulated by sharp transverse ridges. The basal cavity is 
relatively large, circular to ovate, usually with prominent lips. 

Remarks : The Luton form is somewhat more laterally bowed than the specimens 
described by Huddle (loc. cit.). Itis therefore only compared with P. pennatulus. 
Polygnathus bischoffi Rhodes et al. (1969, pp. 184-185, Pl. 13, figs. 8a—11e) is 
a species very close to, if not identical with P. pennatulus. 


Polygnathus cf. symmetricus Branson (Pl. v, Figs. 17-18) 
Cf. Polygnathus symmetrica Branson, 1934, p. 310, Pl. 25, fig. 11; Klapper, 1966, 
p. 21, Pl. 4, figs. 7,9; Pl. 6, figs. 1, 5 (q.v. for full synonymy). 
Figured hypotype: F10901/17. 
Remarks : Although the illustrated specimen has a nearly bilaterally symmetrical 
platform with the anterolateral margins only slightly upturned, its posterior 
end is attenuated. It is therefore intermediate between P. symmetricus and 
P. longiposticus Branson and Mehl (Klapper, 1966, pp. 20-21). Klapper (loc. 


cit.) noted that many specimens in his material also were intermediate between 
these two species. 


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P. symmetricus ranges from to VI strata (Bischoff, 1956) to cw II strata 
(Bischoff and Ziegler, 1956). 


Polygnathus vogesi Ziegler (Pl. v, Figs. 8, 19-22) 

Polygnathus vogest Ziegler, 1962, pp. 94-95, Pl. 11, figs. 5-7. 
Polygnathus styriaca Ziegler mm Fligel and Ziegler, 1957, pp. 47-48, Pl. 1, fig. 11 

(not figs. 12-13 =styriacus). 
Polygnathus cf. styriaca Ziegler, Voges, 1959, p. 294, Pl. 34, figs. 36-41. 
Figured hypotypes: F10901/8, 19-22. 
Description: This species of Polygnathus has a thin, upwardly arched, lanceolate 
platform. The platform is ornamented with two distinct denticulate ridges 
which run from the anterolateral corners to the central part of the platform. 
The anterior margin of the platform is deeply trough-shaped and grooves usually 
are present also posterior to the radiating ridges. Aboral surface has a tiny basal 
cavity mounted on the prominent keel. The lower surface may possess grooves 
complementary to the radiating ridges of the oral surface. 
Remarks : This species closely resembles P. communis carinus Hass, 1959. Voges 
(loc. cit.) has already noted that in this latter form the basal cavity is distinctly 
larger, the radiating ridges are less regular, and the anterior end of the platform 
less sunken. The Australian specimens closely resemble those described by 
Voges. The specimens on which Ziegler (loc. cit.) based the species tend to have 
a somewhat broader platform and a suggestion of nodose ornament on the 
posterior part of the platform. 

In Europe P. vogesi ranges from the Middle costatus Zone (to V ?-to VI) 
(Ziegler, 1962) into cu I strata. 


Polygnathus sp. (Pl. v, Figs. 5-7) 
Figured hypotypes: F10901/5, 7. 


Remarks: This species resembles most closely Polygnathus obliquicostatus Ziegler 
(1962, pp. 92-93, Pl. 11, figs. 8-12 ; Klapper, 1966, p. 22, Pl. 6, figs. 2, 4). In 
its upper surface morphology it differs most notably in that the transverse 
ridges of the inner platform are not so oblique to the carina, and the carina 
extends to the posterior end of the platform. A keel is present throughout the 
length of the unit. 

In the possession of a rostrum, this species, like P. obliquicostatus, resembles 
species of Siphonodella. However, Klapper (1966), in distinguishing Siphono- 
della and Polygnathus, has emphasized that the latter genus has a raised keel 
which runs the length of the platform and is interrupted only by the basal cavity. 
In contrast, the keel of Siphonodella is either absent or is represented by a thin 
sroove which runs some distance posterior to the basal cavity. 

This species, which is abundantly represented in material of Early 
Carboniferous age from Queensland, will be described in detail elsewhere. 


Genus PSEUDOPOLYGNATHUS Branson and Mehl, 1934 
Type species: Pseudopolygnathus primus Branson and Mehl. 


Pseudopolygnathus dentilineatus Branson (Pl. vi, Figs. 23-25) 
Pseudopolygnathus dentilineata Branson, 1934, p. 317, Pl. 26, fig. 22; Bishcoff, 
1957, pp. 50-51, Pl. 4, figs. 29-32, 34; Voges, 1959, pp. 300-301, Pl. 34, 
figs. 49-50; Klapper, 1966, pp. 14-15, Pl. 5, figs. 10-11 (q.v. for full 
Synonymy); van Adrichem Boogaert, 1966, p. 185, Pl. 3, fig. 8; Canis, 
1968, p. 546, Pl. 73, figs. 10, 29-31. 
Pseudopolygnathus dentilineatus Branson, Rhodes et al., 1969, pp. 208-209, Pl. 5, 
figs. 9a-13c; PI. 6, figs. 8a-c. 


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12 LATE DEVONIAN CONODONTS, LUTON FORMATION, N.S.W. 


Pseudopolygnathus vogesi Rhodes et al., 1969, pp. 216-217, Pl. 5, figs. la-c, 3a-c, 
5a-8 ; Druce, 1969, pp. 114-115, Pl. 36, figs. la—7b. 

Figured hypotypes: F10900/23-25. 

Description: The platform is usually narrow and pointed and ornamented with 

coarse nodes or irregular transverse ridges. The right side of the platform extends 

further toward the anterior end than does the left side. Basal cavity large, 

covering the entire width of the platform, and usually has flaring lips. 


Remarks: Rhodes et al. (1969), on the basis of the Carboniferous conodont 
sequence of Britain, postulated three successive independent developments of 
Pseudopolygnathus from species of Spathognathodus with double rows of denticles. 
The first development included forms which they named as P. vogesi and 
considered as homeomorphs of P. dentilineatus. This latter name was restricted 
to a form postulated as originating from Spathognathodus as a later time. No 
morphological characters serve to distinguish these species. Until such time 
as a similar discontinuity in the occurrence of species of Pseudopolygnathus is 
established in N. America (e.g. in the Hannibal Formation in Missouri) it seems 
premature to follow this division. 

Forms of the morphology of P. dentilineatus have a total range from the 
Upper styriacus Zone (to V) to the Z zone of the British Avonian (? ew IT 
beta/gamma). 


Pseudopolygnathus trigonicus Ziegler (Pl. vi, Figs. 16-19) 

Pseudopolygnathus trigonica Ziegler, 1962, pp. 101-102, Pl. 12, figs. 8-13 ; Freyer 

and Zakowa, 1967, pp. 109-110. 
Figured hypotypes: F10900/16-19. 
Description: This species of Pseudopolygnathus has a thin, arched, triangular 
platform and a long high blade. The platform is ornamented with nodes, the 
largest of which form secondary carinae which run to the anterolateral margins 
of the platform. The inner platform usually has a secondary lobe. The aboral 
surface with a central pit and a well-defined posterior keel. Arising from the 
keel are two ridges, one of which runs to the lateral extremity of the outer 
platform and the other to the secondary lobe of the inner platform. 
Remarks: The lateral ridges of the aboral surface delineate a shallow asym- 
metrical depression beneath the anterior part of the platform. This is interpreted 
as the basal cavity, and, as a consequence, the species was referred by Ziegler to 
Pseudopolygnathus. The species cannot be located here without some reservation 
as its upper surface morphology, however, is more akin to that of Ancyrodella. 

Ziegler (1962) originally gave the range of P. trigonicus as Middle to Upper 
costatus Zone. Freyer and Zakowa (loc. cit.) have extended its range to the 
Lower costatus Zone in Poland. 


Genus SPATHOGNATHODUS Branson and Mehl, 1941 
Type species: Ctenognathus murchisoni Pander, 1856. 


Spathognathus aculeatus (Branson and Mehl) (PI. vi, Figs. 12, 20) 
Spathodus aculeatus Branson and Mehl, 1934, pp. 186-187, Pl. 17, figs. 11, 14. 
Spathognathodus aculeatus (Branson and Mehl, Ziegler, 1962, pp. 105-106, Pl. 13, 
figs. 27-36 ; Bouckaert and Ziegler, 1965, Pl. 5, figs. 1-4; Klapper, 1966, 
p. 24, Pl. 6, figs. 15-17; Glenister and Klapper, 1966, pp. 835-836, 
Pl. 95, fig. 11 (q.v. for complete synonymy); van Adrichem Boogaert, 
1966, p. 186, Pl. 3, fig. 19; Freyer and Zakowa, 1967, p. 120, Pl. 1, fig. 12 ; 
Wolska, 1967, p. 425, Pl. 19, figs. 4-8. 

Spathognathodus cf. S. aculeatus (Branson and Mehl), Druce, 1969, p. 124, Pl. 27, 
figs. 5a-e. 


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Figured hypotypes: F10900/12, 20. 


Remarks: The Luton specimens are all broken but are certainly referrable to 
S. aculeatus, as generally interpreted. This form species usually has three to 
five lateral denticles joined to those of the blade by transverse ridges in the 
region of the basal cavity, which is mounted at midlength and is wide and 
circular. 


Recently Rhodes e al. (1969, p. 236) stated that ‘‘ forms described from 
Germany and identified by Ziegler (1961) and other authors may be distinct 
from 8S. aculeatus and are probably a new species”. As no basis for such a 
separation is apparent, here Ziegler’s (1962) and Klapper’s (1966) interpretation 
of the species is followed and the specimens from the Bonaparte Gulf Basin, 
compared by Druce with this species, are included in it. 


Rhodes et al. (op. cit.) also describe a new species, S. plumulus, which differs 
from 8S. aculeatus in the possession of high, crested denticles at the anterior end 
of the unit. 

As interpreted here, the species in Germany ranges from the middle Lower 
costatus Zone to the Middle costatus Zone (i.e. the boundary of to V and VI) to 
to VI alpha (Ziegler, 1962). It has a similar range in Spain (Higgins ef al., 1964 ; 
van Adrichem Bouckaert, 1966), Belgium (Bouckaert and Ziegler, 1965), Poland 
(Freyer and Zakowa, 1967; Wolska, 1967), the Mississippi Valley (Collinson, 
Scott and Rexroad, 1962, pp. 17, 18), the Canning Basin, Western Australia 
(Glenister and Klapper, 1966) and the Bonaparte Gulf Basin, Western Australia 
(Druce, 1969). Canis’s (1968) record of this species from the Bachelor Formation 
of Missouri of cu II alpha age is based on S. antiposicornis Scott. 


Spathognathus crassidentatus (Branson and Mehl) (Pl. vi, Fig. 1) 


Spathodus crassidentatus Branson and Mehl, 1934, p. 276, Pl. 22, fig. 17. 


Spathognathodus crassidentatus (Branson and Mehl), Klapper, 1966, p. 23, Pl. 5, 
figs. 15-17 (q.v. for full synonymy) ; Canis, 1968, pp. 552-553, PI. 74, fig. 29 ; 
Rhodes et al., 1969, pp. 226-227, Pl. 3, figs. la—4b ; Druce, 1969, pp. 127-128, 
Pl. 27, figs. 2a—3b. 

Figured hypotype: F10900/1. 


Description: This is a bladed unit, slightly arched in profile, sloping downwards 
to its posterior end. The anterior end has two enlarged denticles, markedly 
wider and higher than the remainder. The basal cavity extends from somewhat 
in front of midlength to near the posterior end; its greatest width is at the 
rounded anterior margin, whence it tapers to the posterior end. 
Remarks: The Luton specimens agree closely with the paucidenticulate forms 
described by Klapper (loc. cit.) rather than, for example, those illustrated by 
Rhodes et al. (loc. cit.). Most authors, however, have commented on the wide 
range of variation in this species. 

Hitherto, S. crassidentatus has been recorded only from Early Carboniferous 
strata in N. America (Klapper, 1966 ; Canis, 1968), Europe (Rhodes e¢ al., 1969), 
and Australia (Druce, 1969). 


Spathognathodus praelongus Cooper (Pl. v1, Fig. 21) 


Spathognathodus praelongus Cooper in Cooper and Sloss, 1943, p. 175, Pl. 28, 
fig. 14 ; Klapper, 1966, p. 24, Pl. 6, fig. 23 (q.v. for full synonymy). 

Figured hypotype: ¥10900/21. 

Remarks: This is a bladed species of Spathognathodus with the posterior end of 

unit incurved. A slightly enlarged denticle present above the basal cavity, 

which is marked toward the posterior end of unit. Klapper (loc. cit.) gives the 

range of this species as Late Devonian in N. America. 


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Spathognathodus sp. A (Pl. vi, Figs. 11-12) 

Figured hypotypes : F10900/11, 12. 

Remarks: This is a bladed form of Spathognathodus in which the unit tends to 
be incurved. In profile the unit is arched. The basal cavity is symmetrical, 
widest at its anterior end and extending to near the posterior end of the unit. 
All specimens are badly broken, so that the species represented cannot be 
positively identified. It appears to be close to Late Devonian forms identified 
as 8. stabilis (Branson and Mehl) (e.g. Klapper, 1966, PI. 5, fig. 6). 


Spathognathodus sp. B (Pl. vi, Figs. 2-4) 
Figured hypotypes: F10900/2—4. 


Remarks : This is a massive species of Spathognathodus with regular denticulation 
and a prominent asymmetric basal cavity mounted toward the posterior end of 
the unit. Nodes may be present on the upper surface of the lobes of the basal 
cavity. The form apparently bears little relationship to previously described 
Upper Devonian species. 


Genus SYNPRIONIODINA Bassler, 1925 
Type species: Synprioniodina alternata Bassler, 1925. 


Synprioniodina sp. (Pl. vi, Figs. 13, 15) 
Figured hypotypes: F10900/13, 15. 


Remarks: In all of the available specimens the posterior bar is missing, so that 
positive identification is not possible. The cusp and the denticulated anticusp 
compare closely with those of Synprioniodina alternata Bassler (Huddle, 1968, 
pp. 45-46, Pl. 6, figs. 3-5, 12). 


References 


ADRICHEM BooGaERT, H. A. Van, 1967.—Devonian and Lower Carboniferous conodonts of the 
Cantabrian Mountains (Spain) and their stratigraphic application. Leidse geol. Meded., 
39: 129-192. 

BeEerGstrom,, S. M., 1964.—Remarks on some Ordovician conodont faunas from Wales. Acia 
Univ. Lund., 186: 1-66. 

Biscuorr, G., 1957.—Die Conodonten-Stratigraphie der rhenoherzynischen Unterkarbons mit 
Berucksichtigung der Wocklumeria-Stufe und der Devon/Karbon Grenze. <Abh. hess. 
Landesamt. Bodenforsch., 19: 1-64. 

, and ZinGLeR, W., 1956.—Das Alter der “ Urfer Schichten ’ im Marburger Hinterland 
nach Conodonten. Notizbl. hess. Landesamt. Bodenforsch., 84: 138-169. 

BovuckaErt, J., and ZieGLER, W., 1965.—Conodont stratigraphy of the Famennian Stage (Upper 
Devonian) in Belgium. Mem. Serv. géol. Belg., 5: 1—40. 

Branson, EH. B., and Meu, M. G., 1933-1934.—Conodont studies I-IV. Univ. Missouri Stud., 
8 (1-4): 1-349. 

Canis, W. F., 1968.—Conodonts and biostratigraphy of the Lower Mississippian of Missouri. 
J. Paleont., 42): 525—555. 

Coutinson, C. L., Scorr, A. J., and Rexroap, C. B., 1962.—Six charts showing biostratigraphic 
zones and correlations based on conodonts from Devonian and Mississippian rocks of the 
upper Mississippi Valley. Czrc. Illinois State Geol. Surv., 328: 1-32. 

Coopsr, C. L., and Stoss, L. L., 1943.—Conodont fauna and distribution of a Lower Mississippian 
black shale in Montana and Alberta. J. Paleont., 17: 168-176. 

Druce, E. C., 1969.—Devonian and Carboniferous conodonts from the Bonaparte Gulf Basin, 
northern Australia. Bull. Bureau Min. Resour. Geol. Geophys., 98: 1—242. 

Eruineton, R. L., 1965.—Late Devonian and Early Mississippian conodonts from Arizona and 
New Mexico. J. Paleont., 39: 566-589. 

Fuucet, H., and Zinciter, W., 1957.—Die Gliederung des Oberdevons und Unterkarbons am 
Steinberg westlich von Graz mit Conodonten. Mutt. Naturw. Ver. Stevermark, 87: 25-60. 

FREYER, G., and Zakowa, H., 1967.—Famennian Conodonts from borehole Bolechowice [ (in 
the Holy Cross Mts.). Acta Geol. Polonica, 17: 105-137. 

GLENISTER, B. R., and Kiaprrr, G., 1966.—Upper Devonian conodonts from the Canning Basin, 
Western Australia. J. Paleont., 40: 777-842. 

Hass, W. H., 1953.—C and correlati the Barnett Formation of Texas. Prof. Pap. U.S. geol. 
Surv., 243-F: 69-. 


PROCEEDINGS OF THE LINNEAN SocieTy oF NEw SoutH WALES, Vow. 96, Part 1 


G. M. PHILIP AND J. H. JACKSON (13; 


,1956.—Ageonodonts fromon of the Chattanooga Shale and the Murray Formation. 
Ibid., 268: 1—4791. 
, 1959.—Conodonts from the Chappel Limestone of Texas. Jbid., 294-J : 365-399. 

Hetms, J., 1963.—Zur “~ Phylogenese”’ und Taxionomie von Palmatolepis (Conodontida), 
Oberdevon. Geologie Jahrg., 12 (4): 449-485. 

Hiecins, A. C., WAGNER-GENTIS, C. H. T., and WaGner, R. H., 1964.—Basal Carboniferous 
strata in part of northern Leon, NW. Spain: Stratigraphy, conodont and goniatite faunas. 
Bull. Soc. belge Géol. Paléont. Hydrol., 75: 205-247. 

Huppte, J. W., 1958.—Redescription of Upper Devonian conodont genera and species proposed 
by Ulrich and Bassler in 1926. Prof. Pap. U.S. geol. Surv., 578: 1-55. 

JENKINS, T. B. H., 1966.—The Upper Devonian index ammonoid Cheiloceras from New South 
Wales. Palaeont., 9: 458-463. 

, 1968.—Fammenian ammonoids from New South Wales. Jbid., 11 (4): 535-548. 

Kuaprer, G., 1966.—Upper Devonian and Lower Mississippian Conodont zones in Montana, 
Wyoming, and South Dakota. Pal. Contrib. Kansas Univ., 3: 1-48. 

McKetvey, B. C., and Waite, 1964.—Geological Map of New England 1: 100,000, Horton Sheet 
(No. 290), with marginal text. University of New England, Armidale, New South Wales. 

Ment, M. G., and ZiecLER, 1963.—A neotype for Palmatolepis gracilis Branson & Mehl 1934. 
J. Scr. Lab. Denison Univ., 45: 197-206. 

Mounp, M. C., 1968.—Upper Devonian conodonts from southern Alberta. J. Paleont., 42: 
444-524. 

Mutter, K. J., 1956.—Zur Kenntis der Conodonten-Fauna des europaischen Devons, 1. Die 
Gattung Palmatolepis. Abh. senckenb. naturf. Ges., 494: 1-69. 

, and Crarg, D. L., 1967.—EHarly Late Devonian conodonts from the Squaw Bay 
Limestone in Michigan. J. Paleont., 41: 902-919. 

PreppER, A. EH. H., 1968.—The Devonian System of New England, New South Wales, Australia. 
Intern. Symposium of the Devonian System (ed. D. H. Oswald) Calgary, 2: 135-142. 

Puitip, G. M., 1966—Middle Devonian conodonts from the Moore Creek Limestone, northern 
New South Wales. J. Proc. Roy. Soc. N.S.W., 100: 151-161. 

Pickett, J. W., 1960.—A clymeniid from the Wocklumeria zone of New South Wales. Palaeont., 
3: 237-241. 

ReEexroap, C. B., and Scorr, A. J., 1964.—Conodont zones in the Rockford Limestone and the 
lower part of the New Providence Shale (Mississippian) in Indiana. Bull. geol. Surv. Indiana, 
30: 1-54. 

Ruops#s, F. H. T., Austin, R. L., and Drucs, E. C., 1969.—British Avonian (Carboniferous) 
conodont faunas, and their value in local and intercontinental correlation. Bull. Brit. Mus. 
(Nat. Hist.) Geol. Suppl., 5: 1-313. 

Vocss, A., 1959.—Conodonten aus dem Unterkarbon I und II (Gattendorfia- und Pericyclus- 
Stufe) des Saurerlandes. Palaont. Z., 33: 266-314. 

WotskKa, Z., 1967.—Upper Devonian conodonts from the south-west region of the Holy Cross 
Mountains, Poland. Acta Pal. Polonica, 12: 363-456. 

ZIEGLER, W., 1962.—Taxionomie und Phylogenie oberdevonischer Conodonten und ihre strati- 
graphische Bedeutung. Abh. hess. Landesamt. Bodenforsch., 38: 1—166. 


EXPLANATION OF PLATES 


All figures x 40 and specimens registered in the Palaeontological Collection of the University 
of New England. 


PLATE V 


Fig. 1. Bryantodus sp. Lateral view of F10901/1. 
Fig. 2. Hindeodella sp. Inner view of F10901/2. 
Fig. 3. Hibbardella sp. Posterior view of F10901/3. 
Fig. 4. Ozarkodina sp. Lateral view of F10901/4. 


Figs. 5-7. Polygnathus sp. 5, Aboral view of F10901/5. 6, Oral view of same specimen. 
7, Oral view of F10901/7. 


Figs. 8, 19-22. Polygnathus vogesi Ziegler. 8, Lateral view of F10901/8. 19, Oral view of 
F10901/19. 20, Aboral view of F10901/20. 21, Oral view of F10901/21. 22, Oral view 
of F10901/22. 


Fig. 9. Lonchodina sp. Inner view of F10901/9. 

Fig. 10. Ligonodina bicurvata Mound. Inner view of F10901/10. 
Fig. 11. Hindeodella? sp. Inner view of F10901/11. 

Fig. 12. Ozarkodina sp. Lateral view of F10901/12. 


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76 LATE DEVONIAN CONODONTS, LUTON FORMATION, N.S.W. 


Fig. 13. Ozarkodina sp. Lateral view of F10901/13. 

Fig. 14. Lonchodina sp. Inner view of F10901/14. 

Fig. 15. Lonchodina sp. Inner view of F10901/15. 

Fig. 16. Dzplopodella sp. Posterior view of F10901/16. 

Figs. 17-18. Polygnathus cf. symmetricus Branson. 17, Lateral view of F10901/17. 18, Oblique 
oral view of same specimen. 

Figs. 23-27. Polygnathus cf. pennatulus Bassler. 23, Aboral view of F10901/23. 24, Aboral 
view of F10901/24. 25, Oblique oral view of F10901/25. 26, Aboral view of F10901/26. 
27, Oblique oral view of F10901/27. 


PLATE VI 


Fig. 1. Spathognathodus crassidentatus (Branson and Mehl). Lateral view of F10900/1. 

Figs. 2-4. Spathognathodus sp. B. 2, Lateral view of F10900/2. 3, Lateral view of F10900/3. 
4, Oral view of F10900/4. 

Figs. 5-7. Palmatolepis gracilis sigmoidalis Ziegler. 5, Oblique oral view of F10900/5. 
6, Oblique oral view of F10900/6. 7, Oblique oral view of F10900/7. 

Figs. 8-10, 26-29. Palmatolepis gracilis gracilis Branson and Mehl. 8, Oblique oral view of 
F10900/8. 9, Oral view of F10900/9. 10, Oral view of F10900/10. 26, Oral view of 
F10900/26. 27, Aboral view of F10900/27, a broken platform. 28, Oral view of F10900-28, 
a complete specimen with a fully developed platform. 29, Oral view of F10900/29. 

Figs. 11, 22. Spathognathodus sp. A (Branson and Mehl). 11, Lateral view of F10900/11. 
22, Lateral view of F10900/22. 

Figs. 12, 20. Spathognathodus aculeatus (Branson and Mehl). 12, Aboral view of F10900/12. 
20, Oral view of F10900/20. 

Figs. 13, 15. Synprioniodina sp. 13, Inner view of F10900/13. 15, Inner view of F10900/15. 

Fig. 14. Neoprioniodus oligus (Cooper). Inner view of F10900/14. 

Figs. 16-19. Pseudopolygnathus trigonicus Ziegler. 16, Oral view of F10900/16. 17, Lateral 
view of F10900/17. 18, Oral view of F10900/18. 19, Aboral view of F10900/19. 

Fig. 21 Spathognathodus praelongus Cooper Lateral view of F10900/21 

Figs. 23-25. Pseudopolygnathus dentilineatus Branson. 23, Oral view of F10900/23. 24, Oral 
view of F10900/24. 25, Oral view of F10900/25. 


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Proc. LINN. Soc. N.S.W., Vol. 96, Part 1 PLATE V 


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Proc. LINN. Soc. N.S.W., Vol. 96, Part 1 PLATE YI 


A BACTERIAL LEAF SPOT OF ZINNIA IN NEW SOUTH WALES 


A. L. BERTUS 


Biological and Chemical Research Institute, Department of Agriculture, 
Rydalmere, New South Wales, 2116 


AND 
A. C. HAYWARD 


Department of Microbiology, Medical School, University of Queensland, 
Herston, Brisbane, 4006 


(Plates vi, VIII) 


[Accepted for publication 24th March 1971] 


Synopsis 
A bacterial disease of zinnia caused by Xanthomonas nigromaculans f. sp. zinniae is recorded 
for the first time in Australia. Two isolates from material collected at Armidale, N.S.W., were 
pathogenic on inoculation into the zinnia cultivar from which they were originally obtained 
and to eight other cultivars. These cultures were compared with two isolates of the species from 
Africa, including the original culture of Hopkins and Dowson (1949), and were shown to be 
indistinguishable except for some minor differences in carbohydrate utilization. 


INTRODUCTION 


In February 1969, leaves of zinnias growing in the grounds of Duval College, 
University of New England, Armidale, were observed to have irregular blackened 
spots from 1 mm. in diameter to large necrotic areas over 20 mm. across. The 
larger areas seemed to be formed by the coalescing of several small spots. The 
spots had a water-soaked appearance and all lesions were surrounded by yellow 
chlorotic margins (Plate vil, Fig. 1). A massive bacterial ooze was obtained 
from some of the spots on suspension in water. The plants were grown from 
commercial seed of the cultivar Giant Dahlia Flowered Mix. 


The only earlier record of a bacterial leaf disease of zinnia in New South 
Wales is from Cremorne in December, 1948 (Anon., 1949). This specimen is 
filed in the Biology Branch Herbarium at Rydalmere (DAR 4007). The leaves 
show small black spots; it is noted on the herbarium packet that abundant 
ooze was observed on the spots. The organism was not identified. 


A survey of available plant disease check lists of other Australian States 
does not reveal any published record of a bacterial leaf disease of zinnia. 


A bacterial leaf spot and flower disease of zinnia was reported by Hopkins 
and Dowson (1949) from Southern Rhodesia. The causal bacterium was described 
by them and named Xanthomonas nigromaculans f. sp. zinniae. The host 
symptoms described and illustrated closely resemble the symptoms observed 
at Armidale. This disease has also been recorded from Brazil (Robbs, 1954) 
and from Sierra Leone (Deighton, 1956). 


In the following an account is given of the isolation and identification of 
the causative bacterium, which involved pathogenecity tests, cultural and 
physiological observations and electron microscopy. This detailed study was 
undertaken as the descriptions by Hopkins and Dowson (1949) and others are 
inadequate. 


PROCEEDINGS OF THE LINNEAN SociETY OF NEw SoutH Wates, Vou. 96, Part 2 


i.2) 
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BACTERIAL LEAF SPOT OF ZINNIA IN NEW SOUTH WALES 


MATERIALS AND METHODS 


Isolation and cultural and physiological tests: Water-soaked spots were cut 
from leaves and suspended in 3 ml. of sterile distilled water ; after standing for 
two hours at room temperature to allow egress of bacterial ooze, loopfuls of 
suspension were streaked out on 2°% sucrose/peptone agar (Hayward, 1964). 


Two isolates of Xanthomonas nigromaculans f. sp. zinniae, NCPPB 189 
isolated from Zinnia sp. in Southern Rhodesia in 1946 and NCPPB 799 isolated 
from Zinnia elegans in Malawi in 1960, were obtained under licence from the 
National Collection of Plant Pathogenic Bacteria, Plant Pathology Laboratory, 
Hatching Green, Harpenden, Herts., England. This species is the only known 
bacterial pathogen affecting the leaves of Zinnia. The two cultures were 
compared with two isolates obtained from the Armidale collection, accessioned 
as 0365a and 0365b. 


Cultural and physiological tests for identification were carried out as described 
by Allen, Hayward, Halliday and Fulcher (1970). All test media were incubated 
at 28°C. unless otherwise indicated. Yellow pigments from isolates of the 
bacterium were extracted and saponified using the techniques described by Starr 
and Stephens (1964). Solutions of the yellow pigments in petroleum ether 
were examined in a Perkin-Elmer 4000A, UV—VIS recording spectrophotometer. 


Pathogenicity tests : Two isolates of the bacterium were grown on potato dextrose 
agar and a suspension in sterile distilled water was obtained by washing off the 
growth of six one-week-old slopes of each isolate in 50 ml. of water. The 
suspensions were atomized on 18-day-old seedlings from the same lot of seed 
from which the plants showing natural infection were grown. Controls were 
sprayed with sterile distilled water. Each group of plants was isolated in 
separated humid chambers. Plants of eight other cultivars grown from 
commercial seed were inoculated in the same way with one isolate of the bacterium. 


OBSERVATIONS 


Isolation and cultural characteristics: After 48 hours incubation at 28° C., the 
cultures originating from suspensions of ooze from leaf spots showed a glistening 
yellow growth accompanying a mixture of faster-growing bacterial colonies 
which were visible to the naked eye after 24 hours incubation. The slower 
growing yellow colonies were uniform in appearance. Colonies were picked off 
and purified by successive re-streaking. After three days incubation colonies 
of the yellow, slow-growing bacterium were 3-0-—3-5 mm. in diameter, convex, 
circular with an entire edge, amorphous, glistening, viscid and readily emulsified 
in water. On a peptone/yeast extract medium lacking carbohydrate the colonies 
were 1-5—2-0 mm. in diameter after four days incubation at 28° C.; the colonies 
were low convex and markedly less mucoid than on the carbohydrate-containing 
medium. In cultural characteristics, therefore, the bacterium resembled the 
xanthomonads. 


Cultural and physiological tests: The four cultures 0365a, 0365b, NCPPB 189 
and NCPPB 799 were alike in being aerobic, Gram-negative, uniformly stained 
rods, actively motile with a single polar flagellum (Plate vit), which produced 
yellow viscid growth on 2% sucrose/peptone agar. All cultures were oxidative 
in their metabolism of carbohydrates and produced acid under aerobic conditions 
from cellobiose, fructose, galactose, glucose, glycerol, lactose, maltose, sucrose 
and trehalose. NCPPB 189 produced acid from mannitol; the other three 
cultures were negative after 14 days incubation. Arabinose was oxidized 
promptly by NCPPB 189 and slowly by NCPPB 799 and 0365b ; 0365a failed 
to oxidize arabinose. No acid was produced after 14 days incubation from the 


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A. L. BERTUS AND A. C. HAYWARD 83 


following carbohydrates : adonitol, dulcitol, inositol, salicin and sorbitol. The 
following tests were positive for all cultures: aesculin hydrolysis, phosphatase, 
malonate utilization in Leifson’s medium, citrate utilization in Simmons’ medium, 
growth in a peptone/yeast extract medium containing 2°, sodium chloride, and 
in the same medium containing 3% sodium chloride. All cultures were strongly 
lipolytic on Tween 80 agar and gave wide zones of hydrolysis on starch agar and 
on gelatin agar. Twenty-four-hour peptone/yeast extract agar cultures were 
tested for catalase and for oxidase using Kovacs’ method ; all cultures were 
catalase-positive and gave a weak reaction in the Kovacs test, in which there 
was a colour change at 5-15 sec. after application of culture to the reagent- 
impregnated filter paper. Pectate gel medium was progressively softened at 
7-28 days. Raw potato slices 10 mm. thick, heavily inoculated with each of the 
four cultures down a narrow median groove and incubated for 24 hours at 28° C. 
showed a golden to brownish band of growth and rot extending about 5 mm. on 
either side of the groove. The rot did not extend significantly after a further 
24-hour incubation. This confirms the pectolytic activity of the species and 
perhaps indicates that this pathogen is potentially capable of producing a soft 
wet rot in plant tissues. The capacity to rot potato slices is fairly common in 
Xanthomonas. The following tests were uniformly negative: production of 
nitrite from nitrate, production of gas from nitrate, sulphatase, oxidation of 
gluconate, growth in peptone/yeast extract medium containing 6°, sodium 
chloride, and in the same medium containing 10°, sodium chloride, production 
of phenylpyruvic acid fiom phenylalanine, arginine dihydrolase, chitin 
hydrolysis, levan production on 5°% sucrose/peptone agar, production of 3-keto- 
lactoside from lactose. No diffusible pigments were produced on any medium. 
The extracted yellow pigments of cultures 0365a and 0365b were found to have 
the same absorption maxima claimed by Starr and Stephens (1964) to be 
characteristic of a carotenoid alcohol found only in xanthomonads. 


Pathogenicity tests: All inoculated plants of Giant Dahlia Flowered Mix showed 
infection after 14 days. The controls sprayed with water remained healthy. 
The symptoms were similar to those found in natural infection (Plate vm, Fig. 2) 
and similar to published photographs (Hopkins and Dowson, 1949). There was 
no obvious difference between the reactions of the two isolates. The bacterium 
was readily re-isolated from infected plants. 


All plants of the other eight cultivars of zinnia inoculated with one isolate 
of the bacterium became infected and the bacterium was re-isolated in each 
instance. The cultivars were Thumbelina Mixed, Persian Carpet, State Fair 
Tetraploid Mixed, Sprite Mixed, Semi-dwarf Coquette Mixed, Lilliput Mixed, 
Pink Buttons and Linearis. 


DISCUSSION AND CONCLUSIONS 


On symptoms the zinnia disease observed at Armidale appeared similar 
to that originally described by Hopkins and Dowson (1949) from Southern 
Rhodesia and subsequently found in Brazil (Robbs, 1954) and Sierra Leone 
(Deighton, 1956). 

The cultural and physiological properties of the two isolates from zinnia 
leaves from Armidale are wholly consistent with the genus Yanthomonas as 
described by Dye (1962) and others. These cultures closely resemble the two 
cultures from Africa and differ only in the fact that alone of the four cultures 
NCPPB 189 oxidized both mannitol and arabinose. promptly. The differences 
in carbohydrate utilization are trivial and within the range of variation which is 
encountered in other species of the genus Xanthomonas. There is no doubt that 
the two Armidale cultures are correctly identified as Xanthomonas nigromaculans 
(Takimoto) Dowson 1943 f. sp. zinniae Hopkins and Dowson 1949. 


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a 


34 BACTERIAL LEAF SPOT OF ZINNIA IN NEW SOUTH WALES 


ACKNOWLEDGEMENTS 


The support of a research grant to one of us (A.C.H.) from the Rural Credits 
Division of the Reserve Bank of Australia is gratefully acknowledged. We 
thank Mr. J. V. Hardy and Mr. J. A. Fuerst for their help in the preparation of 
the electron micrographs which were taken at the Electron Microscope Centre, 
University of Queensland, St. Lucia; Dr. A. B. Lloyd, University of New 
Hngland, Armidale, for specimens of diseased zinnias and zinnia seed ; and Mr. 
A. Searle, Department of Agriculture, Rydalmere, for the photographs on 
Plate vm. 


References 

ALLEN, R. N., Haywarp, A. C., Hatuipay, W. J., and FuntcuEr, J., 1970.—Bacterial blight of 
Vicia sativa: aetiology of the disease and identification of the pathogen. Aust. J. biol. Sci., 
23: 597-606. 

Anon., 1949.—New plant diseases. Agric. Gaz. N.S.W., 60: 141-142. 

Deieuton, F. C., 1956.—Plant Pathology Section. Rep. Dep. Agric. S. Leone 1954, 27-28. 
(Abstr. in Rev. appl. Mycol., 36: 381, 1957.) 

Dyr, D. W., 1962.—The inadequacy of the usual determinative tests for the identification of 
Xanthomonas spp. N.Z. J. Sci., 5: 393-416. 

Fuerst, J. A., and Haywarp, A. C., 1969.—Surface appendages similar to fimbriae (pili) on 
Pseudomonas species. J. gen. Microbiol., 58: 227-238. 

Haywarp, A. C., 1964.—Characteristics of Pseudomonas solanacearum. J. appl. Bact., 27: 
265-277. 

Hopkins, J. C., and Dowson, W. J., 1949.—A bacterial leaf and flower disease of Zinnia in Southern 
Rhodesia. Trans. Br. mycol. Soc., 32: 252-254. 

Roses, C. F., 1954.—Bacterias fitopatogenicas do Brasil (Phytopathogenic bacteria of Brazil). 
Agronomia, Rio de J., 13: 265-282. (Abstr. in Rev. appl. Mycol., 35: 752, 1956.) 

Srarr, M. P., and STEPHENS, W. L., 1964.—Pigmentation and taxonomy of the genus Xantho- 
monas. uJ. Bact., 87: 293-302. 


EXPLANATION OF PLATES 
PLATE VII 
Fig. 1. Bacterial leaf spot of zmnia. Natural infection, Armidale, N.S.W. x0-6. 
Fig. 2. Bacterial leaf spot of zinnia. Artificial moculation with isolate 0365b. 1-2. 


PLATE VIII 
Fig. 1. Xanthomonas mgromaculans f. sp. zinniae NCPPB 189. 24,540. Preparation 
negatively stained with 0-5% ammonium molybdate. A single polar flagellum is shown. 
Fig. 2. Xanthomonas nigromaculans f. sp. zinniae 0365. 8,180. Preparation negatively 
stained with 0-5°% ammonium molybdate. Single polar flagella are shown. 
[The technique of negative staining used was that described by Fuerst and Hayward (1969). | 


PROCEEDINGS OF THE LINNEAN SocrETY oF New SoutH Watess, Vor. 96, Part 2 


<2) 


Proc. Linn. Soc. N.S.W:, Vol. 96, Part 2 PLATE VII 


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Proc. Linn. Soc. N.S.W., Vol. 96, Part 2 PLATE VIII 


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A NEW SPECIES OF LOPHOPETALUM (CELASTRACEAE), A GENUS 
NEW FOR AUSTRALIA 


N. BYRNES 
Primary Industries Branch, Northern Territory Adminstration 


(Plate Ix) 
[Accepted for publication 24th March 1971] 


Synopsis 
A new species of Lophopetalum is described: Lophopetalum arnhemicum N. Byrnes, from 
Katherine Gorge (14° 20’S., 132° 27’ E.), Northern Territory, Australia. 
The genus is new for Australia. 


INTRODUCTION 


The genus Lophopetalum is widely distributed throughout S.E. Asia and 
the islands to the north of Australia, where it is represented by about 18 species. 
Its discovery in Australia extends the range of the genus, but the species is 
distinct from those of Malaysia. 


Through correspondence with Dr. C. G. G. J. van Steenis, Dr. Ding Hou 
commented that the species was ‘‘ very distinct and interesting, very different 
from the New Guinea species’. Botanical surveys of the sandstone escarpment 
area of Arnhem Land, a previously poorly explored area, are revealing a number 
of new species and extending the range of many known species. This specimen 
was first collected in Katherine Gorge on a preliminary survey during December, 
1966, but its identity as a new species was not realized until 1968. Complete 
material was collected during that year from a tree near the mouth of the gorge. 
The genus Lophopetalum is represented by evergreen trees with opposite or 
decussate leaves and is distinguished from other members of Celastraceae by 
having three-cornered dehiscent capsules with three loculi, each containing several 
seeds and each seed being completely surrounded by a wing. 


Lophopetalum arnhemicum N. Byrnes, sp. nov. 


Arbor 8 m. attingens cortice ramulorum laevi trunci versus basin tessellato 
saepe eroso. Folia opposita petiolo 0-5-1-0cm. longo lamina lanceolata 
6-12 em. longa 1-5-—2-5 cm. lata apice acuta basi anguste cuneata, venatione 
camptodroma. Inflorescentia glabra thyrsiformis usque ad 10cm. longa 
pedunculis ad 1 cm. longis pedicellis 3-5 mm. longis. Flores circiter 1 cm. 
diametro calyce parvo discum vix superante. Petala late ovata usque ad 3 mm. 
longa sine appendiculis. Discus planus vel supre convexus. Stamina brevissima 
antheris ovatis. Pistillum triangulariter pyramidale discum 2 mm. superans. 
Ovarium triloculare ovulis in locula quoque 6—8 capsula triangularis fusiformis 
6-8 cm. longa. Semina plana circiter 3-5 cm. longa 1-0 cm. lata ala inclusa. 
Holotype: Katherine Gorge National Park, N. Byrnes 921; 7 Sep. 1968 (Darwin 
2852). 


Isotypes: K, L, A, CANB, NSW, BRI, PERTH, MEL, NT. 


Paratype: Katherine Gorge National Park, N. Byrnes 1180; 25 Nov. 1968, 
Darwin, K, L, BRI, NSW, MEL. 

A tree to 8m. high. Bark smooth on the branchlets becoming tesselated 
and often eroded on the trunk. Leaves opposite on petioles 0-5-2-5 em. long ; 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH WaALeES. Vou. 96, Part 2 


86 NEW SPECIES OF LOPHOPETALUM (CELASTRACEAE) 


lamina lanceolate, 6-12 em. long, 1-5-2-5em. wide with arched venation, 
acute, narrow cuneate at the base. Inflorescence a thryse, glabrous, up to 10 em. 
long; peduncles to 1 cm. long; pedicels 3-5 mm. long. Buds shortly conical 
or globose to 3mm. diam. Flowers about 1em. diam. Calyx 4mm. diam. 
with depressed triangular lobes under 0-1 mm. long. Petals broadly ovate, 
3mm. long, without appendages. Dise obscurely five-angled, flat or slightly 
convex about 3mm. diam. Stamens half as long as petals with ovate anthers. 
Gynoecium a triangular pyramid exerted about 2mm. beyond the disc; ovary 
trilocular, each loculus containing 6 to 8 ovules. Fruit a triangular spindle- 
shaped capsule 6-8 cm. long, dehiscing loculicidally. Seeds thin, flat, winged, 
3-5x<1-0cm. ineluding the wing. 

The following description of the pollen was prepared by Mr. J. Muller of 
Rijksherbarium. 

Pollen grains in tetrahedral tetrads. Total size 46 (52-3) 55u, individual 
grains spherical, 29 (33-4) 36u. 

Pores circular, 4 (4-8) 6u in diameter, not opposite in tetrad. 

Total wall thickness 2 (3-1) 3-5u. 

Endexine +1-5u thick. 
Columellae distinct, circular, 0-5 in diameter, +1-5u high, arranged in a 
reticulate pattern in single rows. 

Tectum reticulate; lumina isodiametric, 0-5-0-75u in diameter; muri 
0-5y wide, 0-5u high. 


COMMENT 


The pollen of this species belongs to the Lophopetalum B group as defined 
by Ding Hou (Blumea, XVII, I, 1969). It is very similar to that of LZ. rigidum, 
£L. floribundum, L. macranthum, L. javanicum, L. ledermanm, L. torricellense and 
LE. subovatum. 


This species has been found along the Katherine, South Alligator and East 
Alligator River systems generally restricted to the banks of permanent or larger 
seasonal streams in the hilly sandstone areas. It flowers in August and September 
with the fruit maturing to release seeds during December and January. 


Reference 
Drive Hou, 1963.—Celestraceae I. Flora Malesiana, 6: 227-291 (Leyden). 
, 1969.—Pollen of Sarawakodendron (Celastraceae) and some related genera with notes 
on techniques. Blwmea, 17: 97. 


PROCEEDINGS OF THE LINNEAN SoctETY oF New SourH Watuss, Vou. 96, Part 2 


Proc. Linn. Soc. N.S.W., Vol. 96, Part 2 PLATE IX 


NTA 66 
HERBARILM OF NORTHERN TERRITORY 
AUSTRALIA 


No 


Nene Lepho pelalum arnhemicun 
Sp. nov 
loc Ketherine Gorge Netrenel fark 


co WM By-nes G2! Dae 7 Sep (968 


Notes Banks of river ~ Sondy elluvrurn 
- with rocks - SmoH Oree 
TYPE iss a roan a 
D ens ig20 


The holotype of Lophopetalum arnhemicum N. Byrnes. 


- ee ai 

Sree We, iii inti on’) ss 

= 3 P r , ae ii 7 & +i 
: Hy Dia: : : pa ; shy. Srallin® fo | a eri _ ; i 


_ a facigs Per wink Vyas : ab AD ate, 


‘oa 1, iV 


yp ae "seas 
wee 
i iv Aaa 
edad 
on 


Hout 
. 


a ry i) a 


THE FOOD AND FEEDING OF WINTER WHITING (SILLAGO 
MACULATA QUOY AND GAIMARD) IN MORETON BAY 


JOHN L. MACLEAN 
Zoology Department, University of Queensland* 


[Accepted for publication 21st April 1971] 


Synopsis 

A description of the mouth structure and alimentary canal of Sillago maculata is given to 
indicate minor adaptations associated with a bottom feeding carnivore. Adult whiting, those 
longer than 10cm., were found to consume polychaete worms mainly, but also a variety of 
crustaceans, echinoderms, and small fish, as well as pelecypod molluscs, and Glossobalanus. 

The diet of juvenile whiting, which frequent shallower waters, was found to consist largely 
of mysidaceans and amphipods, and smaller quantities of other crustaceans, pycnogonids, 
polychaetes and gastropods. 

In adults there is a change of diet with size. Only fish longer than 15cm. were found to 
consume significant quantities of molluscs; only fish longer than 16-5 cm. consumed Glosso- 
balanus ; and only fish longer than 17 cm. ate brittle stars. No plant material was found in any 
whiting. 


INTRODUCTION 


Sillago maculata is a bottom feeder and forms part of the benthic nekton 
in Moreton Bay. It is a carnivorous fish, although fishermen sometimes assert 
that the fish feed on weed during the spawning season. For most of the year 
adult whiting inhabit parts of the bay which possess a muddy substrate. Some 
time is apparently spent in sandy areas (Roughley, 1916). However, no 
specimens were found in sandy parts of Moreton Bay in the course of this study. 


The following description of the alimentary system of this species indicates 
the minor adaptations correlated with feeding behaviour. 
(1) Dentition. Small villiform teeth are present on both jaws. Fine vomerine 
teeth are also present. Cardiform pharyngeal teeth occur as firm pads, a large 
median pad on the dorsal and two smaller pads on the medio-ventral surface of 
the pharynx, which leads to a short oesophagus. 
(2) Jaws. The jaws are modified to enable the gape of the mouth to be directed 
obliquely downwards. This is accomplished by a membrane which allows 
protrusion of the upper jaw over the lower. A cartilaginous spine attached to 
the premaxilla provides rigidity for the membrane. 
(3) Alimentary canal. The digestive tract is typically carnivorous in proportions 
(Al-Hussaini, 1949). An elongate caeciform stomach is present ; there are four 
pyloric caeca, and a short intestine follows. A nematode parasite frequently 
invades the intestine, and, less commonly, the stomach. 


METHODS AND MATERIAL 


Adult whiting examined for stomach contents were captured by trawl between 
Mud and St. Helena islands in Moreton Bay in March, May, June and September, 
1966. The sampling area was typical of a large part of the bay, having a substrate 
of deep, soft, sandy mud. Depth ranged from two to six fathoms. 

No juvenile whiting were obtained in this sampling area. Juvenile specimens 
were captured in water less than one fathom deep by bait net near the mouth 
of the Brisbane River in June and November, 1966, only. 


* Present address: Fisheries Division, Department of Primary Industry, Canberra, A.C.T., 
2600. 


PROCEEDINGS OF THE LINNEAN Socrety oF New Soutn Wates, Vou. 96, Part 2 


88 FOOD AND FEEDING OF WINTER WHITING IN MORETON BAY 


After capture, the fish were immediately placed in 10°%% formalin. Altogether, 
433 whiting, including 42 juveniles, were examined for stomach contents. The 
size of juvenile fish ranged from 2-8 cm. to 9-6 cm., and the largest fish examined 
were 21cm. Lengths of fish were caudal fork measurements (l.c.f.), that is the 
distance from the tip of the snout to the fork in the caudal fin. 


In this study, only the presence or absence of each type of food organism was 
recorded. More sophisticated analyses, such as volumetric or gravimetric 
techniques (see review by Hynes, 1950), were precluded by the frequently 
disintegrated condition of the food in the stomachs. In many cases, food 
organisms were recognizable by their hardened portions only, such as skulls or 
other bones in fishes, parapodial chaetae in polychaetes, and various appendages 
or exoskeleton in the crustaceans. Stomachs in which no recognizable food 
organisms were found were either empty or contained small amounts of fluid 
mucus. Both these conditions were recorded as empty stomachs. By summation 
of the whole sample, results were converted to percentage occurrence of each 
item in the stomachs. 


No significant variation with time was noted in the sampling period. 
Consequently, analysis of the data has been confined to variations in the diet 
with size of fish. 


TABLE | 
Food Organisms Found in Stomachs of Winter Whiting in Moreton Bay 


Classification Genera or species 


Phylum Annelida 


Class Polychaeta Several species 
Phylum Mollusca 
Class Pelecypoda Several species 
Class Gastropoda One species 
Pyhlum Arthropoda 
Class Pycnogonida One species 
Class Crustacea 
(a) Cirripedia Lepas sp. 
(b) Copepoda (Cyclopoida) One species 
(c) Ostracoda One species 
(d) Malocostraca (Amphipoda) 
Fam. Gammaridae Several species 
Fam. Caprellidae Several species 
Malocostraca (Stomatopoda) Squilla spp. 
5 (Mysidacea) Heteromysis sp. 
s (Decapoda) Penaeus, Metapenaeus, Thenus, 


several species 
Phylum Echinodermata 


Class Ophiuroidea One species 
Class Holothuroidea One species 
Phylum Chordata 
Sub-phylum Hemichordata Glossobalanus sp. 
Sub-phylum Vertebrata 
Class Osteichthyes Several species, including Hyper- 


lophus translucidus McCulloch 


RESULTS 


A list of all identified genera or groups of food organisms found in the 
stomachs of S. maculata is recorded in Table 1. Of the 433 fish examined, 114 
had empty stomachs. 


Initially, the whiting were divided into two broad categories according to 
gonad condition, juvenile and adult, the latter class including all fish longer 
than 10 cm. 


PROCEEDINGS OF THE LINNEAN SociEry or New SoutH Wass, Vou. 96, Part 2 


JOHN L. MACLEAN ~ 39 


The diet of juvenile whiting was found to consist largely of small crustaceans, 
the largest being about 3mm. long. The most prominent were found to be 
mysideans. Amphipods were second in order of abundance, and species of 
Gammaridae and Caprellidae were obtained. The mysids and amphipods were 
the most commonly occurring organisms in the stomachs and when present were 
usually in large quantity. 

The remainder of organisms eaten by juveniles occurred infrequently and 
in small numbers. They include species of Polychaeta, Pycnogonida, Ostracoda, 
Copepoda, and Mollusca. Interestingly, only two of the juvenile whiting 
analysed had empty stomachs when examined. The frequency of occurrence of 
each food organism in the 42 fish examined is shown below in Table 2. 


TABLE 2 


Frequency of Occurrence of Food Organisms in the Stomachs of 
42 Juvenile Whiting 


Number of 


Frequency of 


stomachs occurrence in 
containing stomachs 
Food organisms food containing 
organism food 
(percentage) 
Mysidacea, 5 a 24 60 
Amphipoda .. ve 12 30 
Polychaeta ae 6 15 
Pycnogonida 2 5 
Ostracoda 2 5 
Copepoda 1 2-5 
Gastropoda ; 1 2-5 
Empty stomachs 2 5 
TABLE 3 


Frequency of Occurrence of Food Organisms in the Stomachs of 
391 Adult Whiting 


Frequency of 


Number of Occurrence in 


stomachs stomachs 
Food organism containing containing 
food organism food 
(percentage) 
Polychaeta a ae 173 62 
Pelecypoda oe on 54 19-5 
Ophiuroidea .. oe 29 10-5 
Holothuroidea a 3 1 
Glossobalanus .. a 17 6 
Brachyura a 3 41 15 
Penaeidae a wa 15 5:5 
Squilla .. is fe 8 3 
Other Crustacea at 48 17 
Fish his RY Ae 15 5:5 
Empty stomachs S¢ 112 — 


Polychaete worms are by far the most commonly occurring food organism 
in the stomachs of adult whiting. Bivalve molluscs and crustaceans, especially 
crabs, comprise most of the remaining portion of the diet. Other crustaceans 
eaten in small numbers include prawns, mantis shrimps, barnacles, and the 
shovel nose lobster. Acorn worms (Glossobalanus), small fish, sea cucumbers, 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Wates, Vot. 96, Part 2 


90 FOOD AND FEEDING OF WINTER WHITING IN MORETON BAY 


and brittle stars are also minor dietary items. One specimen only of Gastropoda 
was found. The frequency of occurrence of each food organism in adult whiting 
is Shown in Table 3. ‘‘ Other Crustacea ”’ includes fragments of organisms not 
further recognizable. 


Since there appeared to be some dietary differences between adult whiting 
of different sizes, they were divided into two arbitrary classes, small 
(10-1-15-5 em.) and large (15-6-21-0 cm.), which were compared by ‘‘ t”’ test 
for quantitative differences in dietary organisms. The tests revealed that at 
the 5% level of significance small adult whiting consumed relatively fewer 
molluses, acorn worms and brittle stars, but more crustaceans other than those 
identifiable as crabs, prawns, or mantis shrimps. 


TABLE 4 
Frequency of Occurrence of Specified Food Organisms in Whiting of Various Size Groups 


Number Empty Brittle 
Size range of stomachs stars Crabs Molluscs _- Polychaetes 
(cm.) Fish (Percentage) (Percentage) (Percentage) (Percentage) (Percentage) 
10-1-10-5 10 0 0 20 0 40 
10-6—-11-0 6 0 0 NYG 17 50 
11-1-11-5 14 0 0 i) 7 29 
11-6—-12-0 12 0 0 0 8 42 
12-1-12-5 9 56 0 0 0 50 
12-6-13-0 7 29 0 20 0 60 
13-1-13-5 12 25 0 0 0 78 
13-6-14-0 19 16 0 13 0 50 
14-1-14-5 24 0 0 9 5 45 
14-6-15-0 30 20 0 4 4 46 
15-1-15-5 35 8 0 13 13 53 
15-6—-16-0 17 23 0 8 22 54 
16-1-16-5 14 28 0 0 20 60 
16-6-17-0 16 69 0 20 0 40 
17-1-17°5 15 27 18 9 19 45 
17-6-18-0 25 24 5 0 21 58 
18-1-18-5 37 24 29 14 19 29 
18-6—-19-0 37 27 19 14 19 33 
19-1-19-5 23 35 13 20 7 47 
19-6—20-0 16 69 0 0 tt) 80 
20-1-20-5 13 8 17 25 25 25 
20-6—-21-0 6 7] 20 20 0 40 


A more detailed division of the adult whiting into 0-5 cm. size groups showed 
that acorn worms were found in fish longer than 16-5 cm. only, and brittle stars 
in fish longer than 17-0cm. Molluscs became “ regular ’’ food items only in 
whiting longer than about 15cm. The occurrence of “‘ other Crustacea ” became 
insignificant in fish longer than 16 cm. Table 4 shows the percentages of whiting 
in the various size groups whose stomachs contained one or more food items. 


DISCUSSION 


The dietary differences between juvenile and adult whiting are significant 
with regard to feeding behaviour. The minute crustaceans and pycnogonids 
which comprise the bulk of the diet of juvenile whiting are planktonic or cling 
to weeds. Bottom-living fauna, the small polychaete worms, are minor food 
items of juvenile fish. Thus, juvenile whiting are basically not bottom feeders. 
On the other hand, the adult whiting diet consists mainly of comparatively large 
benthic organisms, especially polychaetes and pelecypods. The change of diet 
apparently occurs during the movement of juvenile whiting from the shallows 
into deeper waters of the bay. 


PROCEEDINGS OF THE LINNEAN Soctety or NEw SoutrH Watss, Vou. 96, Part 2 


JOHN L. MACLEAN 91 


The selection by larger adult whiting of acorn worms, brittle stars and 
molluses is difficult to explain. The species of brittle star consumed is one of 
the most commonly occurring invertebrate organisms in Moreton Bay, although 
apparently undescribed (Stephenson, personal communication). Pelecypoda 
possess hard shells which may deter smaller fish from eating them. The 
pharyngeal teeth, by which hard particles such as shells are broken up, probably 
strengthen with age. 


The finding of less identifiable remains of crustaceans (‘‘ other Crustacea ’’) 
in large fish may also be correlated with a hardening or strengthening of teeth 
and jaws, such that less time is required to grind the food to a size which allows 
efficient digestion. If so, the frequency of occurrence of recognizable portions 
of the various crustacean food organisms may be correspondingly less. On the 
other hand, larger fish may simply tend to ignore most crustaceans, except 
crabs, as food. 


TABLE 4—Continued 


Number Acorn Mantis Other 
Size range of Worms Prawns Shrimps Crustacea Fish 
(cm.) Fish (Percentage) (Percentage) (Percentage) (Percentage) (Percentage) 
10-1-10-5 10 0 0 0 40 0 
10-6—-11-0 6 0 0 0) 16 0 
11-1-11-5 14 0 0 0 35 29 
11-6—-12-0 12 0 17 8 17 8 
12-1-12-5 9 0 0 0 50 0 
12-6-13-0 7 0 0 0 20 0 
13-1-13-5 12 0 0 0 22 0 
13-6—-14-0 19 0 18 0 19 0 
14-1-14-5 24 0 5 9 27 0 
14-6-15-0 30 0 4 8 26 8 
15-1-15-5 35 0 0 3 12 6 
15-6-16-0 iz 0 8 0 8 0 
16-1-16-5 14 0 20 0 0 0 
16-6-17-0 16 20 0 0 20 0 
17-1-17-5 15 0 9 ) 0 0 
17-6-18-0 25 5 5 0 6 0 
18-1-18-5 37 3 3 0 0 3 
18-6-19-0 37 3 4 4 0 4 
19-1-19-5 23 a 0 0 6 0 
19-6—20-0 16 0 20 0 0 0 
20-1-20-5 13 0 0 0 0 8 
20-6-21-0 6 0 0 0 20 0 


The problem of relative importance of the various food organisms of 
S. maculata is pertinent to this study. Since volumetric and gravimetric methods 
were not applicable, the biomass of each item cannot be determined. In adult 
whiting the most frequently occurring food organisms, polychaetes, were also the 
largest, so that there is no question as to the main source of their metabolic require- 
ments. The problem is significantin juvenile whiting, however. Food organisms of 
juveniles fall into two classes according to size: the minute planktonic forms 
and the somewhat larger polychaetes. The numbers of the former, when they 
occurred in the stomachs of the juvenile whiting, usually exceeded polychaetes 
at least sixfold, one worm fragment only being normally found in the stomach 
of a single juvenile fish. On the other hand, the polychaete pieces eaten were 
larger than the other food organisms, and thus polychaetes may approximate, 
or even exceed, the biomass of other foods eaten by the juvenile whiting. 


Food preferences in fish are well known (Blegvad, 1930; Ivlev, 1961), 
and it is apparent that some form of food selection is exhibited by whiting. 


PROCEEDINGS OF THE LINNEAN Society oF New Sours Waues, Vou. 96, Part 2 


92 FOOD AND FEEDING OF WINTER WHITING IN MORETON BAY 


Knowledge of the relative abundances of the benthic fauna in the whiting feeding 
grounds is required to ascertain the extent of this selection. It is hoped that the 
degree of preferences for different organisms by whiting in Moreton Bay can be 
established in the future, since Stephenson (personal communication) is currently 
investigating the abundance and distribution of benthic organisms there. 


TAINTING 


Fishermen frequently claim that adult whiting eat an iodine-rich weed during 
the spawning season. The basis of this claim is a characteristic odour, reminiscent 
of iodine, possessed by some mature specimens. Its origin was not discovered 
in the present study, and although some fish were found to possess this odour,. 
none contained plant food of any type in their stomachs or intestines. The 
occurrence of the taint is of no commercial importance and it has not been 
sufficiently widespread to warrant investigation by fisheries authorities. 


References 


Au-Hussaini, A. H., 1949.—COn the functional morphology of the alimentary tract of some fishes 
in relation to differences in their feeding habits. Pt.1. Quart. J. Micr. Sci., 90: 109-139. 

BuLeecvaD, H., 1930.—Quantitative investigations of bottom invertebrates in the Kattegat with 
special reference to the plaice food. Rep. Dan. Biol. Stat., 36: 3-55. 

Hynes, H. B. N., 1950.—The food of freshwater sticklebacks (Gasterosteus aculeatus and Pygosteus- 
pungitius), with a review of methods used in studies of the food of fishes. J. Anim. Ecol.. 


19: 36-58. 

Ivurv, V. S8., 1961.—Haxperimental Ecology of the Feeding of Fishes. Yale University Press, New 
Haven. 

Roucuuey, T. C., 1916.—Fishes of Australia and their technology. Sydney Tech. Mus., Educ. 
Ser., 21. 


PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SoutH WALES, VoL. 96, Part 2 


THE EARLY HISTORY AND RELATIONSHIPS OF THE NEW GUINEA 
HIGHLAND DOG (CANIS HALLISTROMTI) 


ELLIS TROUGHTON* 


[Accepted for publication 17th February 1971) 


Synopsis 

There are factual reports of a “ barkless ” dog from east coastal New Guinea from as early as 
1606. Subsequent accounts indicate that this distinctive small dog was forced to the seclusion 
of a mountainous habitat by a combination of hostile circumstances: such as the eating of its 
progeny by natives, as reported from Mt. Giluwee, coupled with its lack of pig-hunting ability. 
This upland migration evidently occurred prior to hybridization with any introduced breed of 
domesticated canine, as indicated by the fact that the discovery of a small wild dog at about 
7,000 ft. on Mount Scratchley in 1897 was considered a faunal novelty by the Administration, 
according to the Annual Report of the Lieut. Governor, Sir William Macgregor (1898). The 
term “ feral” as applied to the New Guinea Dog by some authors is rejected as being irrelevant 
regarding the issue of its specific relationship. 


In Diego de Prado’s Relacion of Torres’ voyage through the Strait in 1606, 
as translated from the Spanish in H. N. Stevens (1930) New Light on the Discovery 
of Australia (p. 141) Prado wrote from the Island of San Facundo [S.E. Papua] 
in 1606: ‘‘ We found small dumb dogs that neither bark nor howl, and do not 
ery out even if beaten with sticks.’ One is indebted to an old colleague, Gilbert 
Whitley, for this most significant reference, as well as several following made 
available from his bibliographical researches. 


In an anonymous article on “* Australasian Islands ”’ in the Colonial Magazine, 
Vol. 7, 1842, p. 62, it is stated that ‘“‘ The bush-dog (Canis Australiae familiaris) 
is an inhabitant of the country Papua: ... these dogs do not bark, but raise a 
howling whine, on being disturbed from their repose. . . Dogs form a portion of 
their native animal food.” In 1886 in the Voyage of the Bonito: an account 
of the Fly River Expedition to New Guinea, in the Mitchell Library, William 
Bauerlen (pp. 28-29) refers to ‘‘ Native dogs which cannot bark, on the Upper 
Fly or Strickland ”’ rivers. 


Alexander Morton, in ‘‘ Notes on a trip to the islands of Torres Strait and 
the south-east coast of New Guinea in Proc. Roy. Geograph. Soc. Austr., N.S.W. 
and Vict. Branches, Vol. 1, 1883, p. 69, states that ‘‘ Motu natives of the Port 
Moresby area in 1877 in their legends attributed the gift of speech to the Dogs”. 
However, Miklouho-Maclay (1881), comparing the ‘‘ Cireumvolutions of the 
Cerebrum ”’ of the Dingo and Papuan Dog, concluded that the superior complexity 
of the Dingo brain indicated differences in modes of life of the respective canines. 
He wrote: ‘‘ The dingo depends upon his skill to provide himself with sufficient 
food by hunting, which requires often the full use of his intellect.” The New 
Guinea Dog “is very timid, and howls instead of barking. . . is used as food by 
the natives of the Maclay Coast; his flesh however, when baked or boiled is 
dark and dry, and is generally regarded by the natives as inferior to that of pig. 
Although he is sometimes fed by his master and gets with the pigs, all the 
remnants of the meals, he is often to be seen on reefs at low water, feeding on 
crabs and small fishes, etc.” 

‘* During the night, the dog in company with the pig takes over the business 
of the ‘ Nightman ’ and the ‘ scavenger ’ of the Papuan villages. In only a very 


* Contributed to the Anthropology Section at the 42nd Congress of A.N.Z.A.A.S. at Port 
Moresby, Papua, on 20th August, 1970. 


PROCEEDINGS OF THE LINNEAN Socrety or New SoutH Wates, Vot. 96, Part 2 


94 EARLY HISTORY AND RELATIONSHIPS OF NEW GUINEA HIGHLAND DOG 


few instances have I seen the natives of the Maclay Coast using their dogs to 
assist them in their hunting the wild pig, the dorcopsis wallaby and the 
cassowary ... 1 think it is not only too timid, but too lazy and stupid an animal 
for such an occupation.” 


In his book on ‘‘ The Orokaiva Society ’’, F. E. Williams (1930) writes of 
‘“ The food quest: Dogs are treated affectionately enough by their owners... 
However, the dog does not compare with the pig in the Orokaiva estimation. 
It is always less a pet than a servant, and is ultimately destined for the pot. 
In slaughtering it is seized by the hind legs and its body dashed against a tree, 
a method which seems brutal but is instantly effective! Many specimens are 
mangy diseased. They are proverbially susceptible to the cold and are commonly 
seen crouching in the ashes of a hardly expired fire. Indeed the dog is said to 
have a special prerogative over the fire by virtue of the legendary fact that he 
first brought fire to the Orokaiva.’’ A footnote adds: ‘ The native dog of the 
Purari Delta is no doubt a degenerate of the same species. From the canine 
point of view the Delta must be a very unfortunate environment.”’ 


In his Annual Report on British New Guinea (1897-1898) Macgregor, referring 
to an extensive patrol of the Mount Scratchley region at 5,000—7,000 ft., wrote 
that where there is such an ample variety of food ‘‘ Animals are rare. The wild 
dog, an occasional wallaby, a stray tree-kangaroo, and a long-snouted animal 
not yet obtained, form the bulk of the quadrupeds.”’ Of the two dog specimens 
subsequently lodged in the Queensland Museum, it is recorded that they were 
obtained from natives, possibly through the interest of white prospectors in the 
area, resulting from the Administrative patrol. 


The first specimen, evidently received from Sir Wm. Macgregor, was described 
and figured by De Vis (1911) of the Queensland Museum. His generalized 
description accounts for the unduly dingo-like illustration in stating that the 
mounting was from a skin long macerating in an entirely boneless state. My 
examination of the two skins proved his colour description to be rather misleading 
in giving the overall colouring as ‘‘ black and white, the black predominant ”’. 


When reviewing the specific status of the dingo, Wood Jones (1925) stated 
that ‘‘ examination of a series of Dingo skulls can leave no doubt as to the true 
affinities of the Australian animal... falls into line with all other races of 
domestic dogs in being of the true northern wolf type. Moreover, in the large 
size of its last upper premolar, or carnassial tooth, it approaches nearer to the 
ancestral type than do the other races of dogs of which I am able to obtain 
specimens or records.” 


With special reference to the relative proportions of this carnassial tooth, 
Heber Longman (1928), in ‘‘ Notes on the Dingo, the Indian Wild Dog, and a 
Papuan Dog’, speculated upon a common origin for the Dingo and Indian 
Wild Dog. He regarded the two specimens of Papuan Dog in the Queensland 
Museum as representing ‘‘ a small breed of true dogs, possibly not truly feral 
or autochthonous... apparently very closely related to the Dingo’. However, 
Longman noted with interest that ‘‘ this race agrees (in carnassial proportions) 
with the Indian Wild Dog (21-5 mm.), the Gray Wolf (25), and average of ten 
dingoes (20) in the relatively large proportion of the carnassial tooth, which is 
more than 10% of the basicondylor length of the skull’’. The relative pro- 
portions of upper carnassial to basi-condylor length in the two Papuan Dogs is 
given respectively by Longman as 15-5 to 149 and 16 to 146 mm. 


In his paper on ‘‘ The cranial characters of the Papuan Dog ’’, Wood Jones 
(1929) refers to Longman’s paper and to borrowing the two skulls of Papuan Dog 
for examination and description. On any doubts concerning their feral nature 
he stated ‘‘ no opinion can be expressed... but it is abundantly evident that 
both skulls are of dogs of precisely the same type. ..and any suspicion of their 
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a 


ELLIS TROUGHTON 95 


being of impure breed is negatived by this remarkable identity. It is certain 
that these two skulls are of representatives of the same race and that this race 
is a primitive one.” 

‘“‘ The skulls are of those of small dogs of the terrier type, with fairly elongated 
noses and well developed muscular crests and ridges. Their most outstanding 
characters are: (a) The form of the supraorbital processes, which are pointed, 
flat to concave on the upper surface, and singularly vulpine in character : 
(b) The sagittal crest, which is formed by a coalescence of the temporal ridges in 
the region of the coronary suture in both specimens, and which is extremely 
prominent in its caudal extremity where it joins the well marked nuchal crest : 
(c) The relatively large size of the upper carnassial tooth, the maximum length of 
which is nearly ten percent of the condylobasal length: (d) The backward 
prolongation of the palate behind the last molar tooth, a very well marked 
distinction of both skulls’ (Text-figs. 1 and 2). 


Text-fig. 1. After Wood Jones, American Journal of Mammology, Vol. 10. 1929. Illustrating 
some of the cranial features referred to in Wood Jones paper. 


Text-fig 2. After Wood Jones, American Journal of Mammology, Vol. 10, 1929. Mlustrating 
some of the cranial features referred to in Wood Jones paper. 


Wood Jones concludes ‘‘ Although these two skulls constitute the only 
material available for study of this dog, they probably afford definite evidence 
that the Papuan feral dog is a very definite race, possessing a relatively large 
upper carnassial tooth typical of primitive canine breeds, and differing widely 
in its characters from the dogs of certain other Pacific islands. It is much to be 
desired that more material should be secured before the crossing by the white 
man’s dogs has rendered the breed too hybridized for the purposes of study.”’ 


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96 EARLY HISTORY AND RELATIONSHIPS OF NEW GUINEA HIGHLAND DOG 


In this regard a ‘‘ Note on New Guinea Dogs ”’ (1952) by the late Dr. G. H. H. 
Tate, then a Curator of Mammals at the American Museum of Natural History, 
becomes of special significance. Briefly reviewing the above contributions on 
the Papuan Dog, Tate records the Archbold collecting of five specimens along the 
Idenburg River, on the northern slopes of the central ranges in West Irian. 
Tate records that ‘‘ In every instance the carnassial length is more than 10 per 
cent of the condylo-basal length in the Idenburg River specimens’. After 
some reference to range of colour, and absence of dew claws in both dingo and 
Papuan Dog, Tate refers to Wood Jones’ conviction that the latter was a primitive 
race. 

Thus we have a very considerable extension of the highland range of a small 
dog of the same type in major cranial characters, from the opposite coast and 
beyond the border of the Territory of New Guinea. While personally collecting 
for the Australian Museum around Mount Hagen in 1954, one had hoped to obtain 
specimens from the relatively intermediate locality of Mount Giluwee, where our 
host-mentor, Mr. N. E. P. Blood, had observed the dogs around villages at about 
8,000 ft. on the 14,346 ft. mountain. Unfortunately, or possibly otherwise for 
myself, Mr. Blood of the Agricultural Station at Mt. Hagen found it unwise from 
an Administrative view to commit one to a 4-5 days’ “ walk ’’ accompanied 
only by a native constable and “ cook-boy”’. A pair of the dogs was later 
obtained by an Assistant District Officer, which, because of their heavy toll on 
poultry around his Station, had to be sacrificed to the villagers. 


Consequently, when Sir Edward Hallstrom arranged to receive a pair of 
the dogs at Taronga Park Zoo, one had no hesitation in naming them specifically 
for the President of the Trust, and benefactor of the Australian Museum. The 
living animals, Canis hallstromi (1957) were so named to offset the inevitable 
claims of ‘‘ Zoo hybridization”’’. The male holotype No. M.8502 and female 
allotype No. M.8917 in the Australian Museum, came from the then uncontrolled 
country in the remote Lavani Valley in the Southern Highlands District of Papua. 
The relative carnassial to condylobasal lengths in the holotype are 15-1 to 
151mm., and 15-5 to 149 in the allotype; and are therefore within the 
characteristic 10°, ratio according to Wood Jones. 


Actually, E. P. Ramsay (1879) had named a dog from the south-east coast 
of Papua Canis familiaris var. papuensis, mainly on the reported inability to 
bark. The specimen was received from the collectors, Alexander Morton and 
Andrew Goldie, and “ unfortunately arrived in a very bad state ” and evidently 
was destroyed. Whether Ramsay’s name be accepted, or rejected as a nomen 
nudum, it has since been overlooked by all authors with the exception of Miklouho- 
Maclay (December, 1881). Even Tate (1952), while listing Maclay’s paper in 
his note on the ‘*“‘ Papuan Dogs” did not refer to Ramsay’s name papuensis, 
though quoting De Vis’ statement that the Mount Scratchley dogs ‘‘ might well 
be feral ”’. 

Throughout this review the specific relationships of the New Guinea Dog 
have been obscured by the irrelevant use of the term ‘‘ feral’, a taxonomic if 
not phylogenetic misnomer, without due allowance for the effect of prolonged 
restriction to a tropical environment. However, one may hope that specific 
naming of the New Guinea Dog will have served its purpose in bringing con- 
sideration of the actual status of the small canid, and its historical association 
with man, before world students of both Mammology and Anthropology, according 
with the expressed aims of Professor Wood Jones (1929). 


As evidence of such interest, a Mrs. I1 Ohman, of Hagford, has written of a 
Swedish research team’s great interest in the New Guinea Dog and of their great 
anxiety that purity of the breed should be assured in captivity ; referring also 
to a paper by a Dr. Von Rosen ‘‘ attempting a definition as a species...as 
highly interesting and decidedly more primitive than the dingo, in general type ”’. 


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ELLIS TROUGHTON 97 


At the request of its author, Mrs. Ohman kindly mailed me a copy of the paper 
by Dr. Wolfhart Schultz (1969) on ‘‘ Zur Kenntnis des Hallstromhundes (Canis 
hallstromi Troughton 1957) ”’. 


The summary in English, after reviewing several domestic breeds of dog, 
refers to an examination of ‘‘ descendants of the original two Papuan Dogs, bred 
in Kiel University Institut”. In his statement that a wide range in coat 
coloration in the two specimens does not prove them ‘‘ to be members of a 
genetically uniform population ’’, Dr. Schultz apparently overlooks the fact that 
the litters of the original pair were entirely black at birth, as are those in San 
Diego Zoo and elsewhere, providing a natural basis for adult variation. 


In view of the known fact that the New Guinea Dog was evidently well 
established in its present form prior to 1606, one finds it impossible to agree 
with Dr. Schultz’ conclusion ‘“‘ that the Hallstrom-dogs are dogs returned to a 
wild state, which are in close relation to the Dingo”’. A conclusion apparently 
overlooking the factors of time, isolation, and the extent of zoo-geographical 
barriers. 


In a note on ‘*‘ The Dog of the Tengger”’, Dr. F. A. Jentink (1896) writes 
of a peculiar kind of dogs living on Mount Tengger, the highest in East Java, 
which had been named Canis familiaris var. tenggerana by Dr. Kohlbrugge 
and which ‘‘ could not live without mountains, they died not by ‘ Heimweh’, 
but by the heat of the lower regions, where they participated with different 
diseases, where they pined away and languished’’. Two complete skeletons 
of the wild Tenggerese dog were presented to the Leyden Museum by Dr. 
Kohlbrugge. Photos of the skull show the relatively large proportions of the 
carnassial tooth. This account would seem to support my previous assumption 
regarding an early upland migration of the New Guinea Dog to its present 
highland habitat. 


Actually, it is my assumption that the ancestral stock of the New Guinea 
Dog arrived by an Asian route and became established in their present form, 
prior to the deployment of the Pacific Island Dog of Hawaiian origin. A wider 
study of the relevant material may provide, according to Wood Jones, ‘‘ a general 
factor in solving the problems of racial movements and racial origins of man and 
the mammals which may have accompanied or preceded him’. At Mount 
Hagen, in 1954, while Mr. N. H. P. Blood was assembling carriers for our several 
patrols, one was impressed by the yodelling calls used by the natives, simulating 
the vocal sounds of the ‘‘ voiceless > New Guinea Dog, and reminding one of the 
Motu legend regarding the gift of speech. Suggestive also of an age-old 
association with man, though not of course the order of it. 


CONCLUSION 


A Papuan region covering the obvious biological unity of the mountainous 
region of Papua with the Atherton Tableland of N.E. Queensland, was proposed 
by Charles Hedley (1894). The types of vegetation, reptiles, birds, and marsupials 
were such, according to Hedley, that any traveller in the heart of the Atherton 
rainforest ‘‘ could scarcely answer, from the surroundings, whether he were 
in New Guinea or Australia ”’. 


Therefore, according with the recent discovery of skeletal remains of the 
Thylacine or ‘‘ marsupial wolf ’’ at 5,000 ft. (1,525 m.) in the Central Highlands 
of New Guinea, and doubtless its past occurrence on the Atherton Tableland, 
in view of persistent reports of a large striped marsupial having inhabited the 
Tableland, it is now postulated that the New Guinea Highland Dog had a similar 
prehistoric co-distribution, and became ancestral to the wild dog or Dingo of 
the Australian mainland, which provided an ample field for the development 
of the Dingo’s hunting resources and physical growth. 


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98 EARLY HISTORY AND RELATIONSHIPS OF NEW GUINEA HIGHLAND DOG 


References 


ANONYMOUS ARTICLE, 1842.—Australasian Islands. Colonial Magazine, 7: 62. 

BAUERLEN, WILLIAM, 1886.—The Voyage of the Bonito: An Account of the Fly River Expedition 
to New Guinea, pp. 28-29. 

Dz Vis, C. W., 1911.—A dog from British New Guinea. Ann. Qld. Mus., 10: 19-20, Pl. i. 

HEDLEY, CHARLES, 1894.—The faunal regions of Australia. Rept. Austr. Assoc. Adv. Science, 
5: 444. 

JENTINE, Dr. F. A., 1896.—The Dog of the Tengger. Notes Leyden Mus., 18: 217-220. 

Jones, F. Woop, 1925.—The mammals of South Australia. Handbook, 3: 151-157. 

, 1929. The cranial characters of the Papuan dog. Amer. Journ. Mammology, 10 (4) : 

329-333, figs. 1-2. 

Loneman, HEBER A., 1928.—Notes on the dingo, the Indian wild dog, and a Papuan dog. Mem. 
Qld. Mus., 9 (2): 151-157. 

MacereGor, Sir Wmn., 1898.—Annual Report on British New Guinea, 1897-1898, p. 27. 

Mactay, N. DE Mixiouno-, 1881.—On the circumvolutions of the cerebrum of Canis dingo. 
Proc. Linn. Soc. N.S.W., 6 (3): 624-626. 

Morton, ALEXANDER, 1883.—Notes on a trip to the islands of Torres Straits and the S.E. coast 
of New Guinea. Proc. Roy. Geogr. Soc. Austr., N.S.W. and Vict. Branches, i: 69. 

Prapo, Dreco pE, 1606.—Relacion of Torres’ voyages through the Strait, as translated from the 
Spanish by N. H. Stevens (1930). In New Light on the Discovery of Australia, p. 141. 

Ramsay, E. P., 1879.—Proc. Linn. Soc. N.S.W., 3 (3): 242. 

Rosen, Dr. Von, 1953.—Journal of the Swedish Kennel Club. 

Scouuz, Dr. Von Wo.urHart, 1969.—Zur Kenntnis des Hallstromhundes (Canis hallstromi 
Troughton 1957). Sonderdruck aus Zool. Anz., 183 (1-2): 47-72. 

Tats, G. H. H., 1952.—Archbold Exped., No. 66, Mammals of Cape York Peninsula (note on New 
Guinea dogs). Bull. Amer. Mus. Nat. Hist., 98 (7): 613-614. 

Troucuton, Exiis, 1957.—A new native dog from the Papuan highlands. Proc. Roy. Zool. Soc. 
N.S. Wales, 1955-1956: 93-94. 

, 1967.—Furred Animals of Australia. 9th Edition, pp. 48—52. 
Wiutiams, F. E., 1930.—The Orokaiva Society, p. 62. 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wates, Vou. 96, Part 2 


AN UNDESCRIBED SPECIES OF USTILAGO ON CY PERUS LUCIDUS R.Br. 
IN AUSTRALIA 


WITH COMMENTS ON ROBERT BROWN’S TYPE COLLECTION OF C. lucidus 


J. WALKER 


Biology Branch, Biological and Chemical Research Institute, 
N.S.W. Department of Agriculture, Rydalmere, N.S.W. 


(Plates x, x1) 
[Accepted for publication 17th February 1971] 


Synopsis 

An ovary smut of Cyperus lucidus R. Br. found in New South Wales is described as Ustilago 
cypert-lucidt n. sp. It was also found in one collection of C. lucidus from Queensland and one 
from Victoria. Hxamination of two portions of Robert Brown’s original gathering of C. lucidus, 
made in N.S.W. in 1802-1805, showed that the smut was present in the portion filed in the 
Herbarium, Royal Botanic Gardens, Kew, but not in the lectotype specimen in the British Museum 
(Natural History). The relationship of U. cyperi-lucidi to other species with pale spores, and 
especially to Ustilago spp. on Cyperaceae and Juncaceae, is discussed. 


INTRODUCTION 
During an examination of collections of Cyperus spp. at the National 
Herbarium of New South Wales, Mr. O. D. Evans found abnormal nuts in some 
collections of Cyperus lucidus R. Br. Detailed examination showed them to be 
filled with a vellowish nowder comnosed of snores of an hitherto unrecognized 


Mine 
‘ions 
CORRIGENDA 

P. 100.—For Plate IX, read Plate X. 

For Plate X, read Plate XT. 
P. 102.—For Plate X, read Plate XT. itiae 
P. 103.—For Plate X, read Plate XI. eat 
P. 104.—For Plate X, read Plate XI. pitis 
P. 107.—For Plate IX, read Plate X. lina, 
For Plate X, read Plate XI. lata, 
oris. 
S et 


primitus 10-15 in fasciculis laxe adherentes, demum singulatim, interdum cum 
frustillis affixis hypharum, (15) 16-22 (25) um diam., spora gigantea rara ad 
27 wm diam., vel sporae ovales 17-23 (25)x12-17 ym (minus quam 10% 
sporarum), pallide viridi-flavae usque pallide flavobrunneae, pariete 1-5—2-0 um 
lato, episporio subtiliter reticulato vel imperfecte reticulato, reticulum 1 -5-2-0 um 
altum, areolis 1-5—2-0 (2-5) um diam. Sporae in solano glucoso agaro ad 15—21° C. 
in diem per promycelium 3-septatum 26-33 x 5-9 um germinantes et sporidia 
ovalia 7-10 x 4-6 um a omni cellula formata. 

Habitat: In acheniis Cyperi lucidi in Nova Wallia Australi, Victoria et Terra 
Reginae, Australia. 


Holotypus: In pasture, property of R. Skellern, Wyndham, west of Pambula, 
N.S.W., 11.vii.1969, J. Hindle, DAR 17587a. 


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100 UNDESCRIBED SPECIES OF USTILAGO CYPERUS LUCIDUS R.Br. 


U. cyperi-lucidi forms its sori within the nuts and in most collections all 
nuts in an infected inflorescence are smutted. Infected inflorescences are very 
difficult to detect except by a detailed examination of the florets, when the 
glumes are seen to be slightly more spread than in healthy florets. They are 
pushed open by the swollen diseased nuts between them (Plate rx, Fig. A). 
Sori are grey, elliptical to broadly obovate, and roughly the same length as 
normal nuts but somewhat wider (Plate 1x, Fig. C). They measure 1-5-2-0 mm. 
long and 1-1-5 mm. wide; nuts are roughly 2 mm. long and 0-5—0-8 mm. wide. 
In section, sori have a more rounded outline than the three-angled outline of 
normal nuts. 


The sori are covered by the unaltered wall of the nut, with no sign of a fungal 
sheath. At maturity, they split open, often along one side (the central sorus in 
Plate 1x, Fig. C) to expose the pale yellow (Munsell (1967) 7-5Y 9/4-6 ; Ridgeway 
(1912) Naphthalene Yellow) to pale yellow brown (Munsell 2-5Y 9/4-6 ; Ridgeway 
Warm Buff) spore mass. Some sori have a darker, pale reddish brown (Munsell 
2-5YR 7/10; Ridgeway Ochraceous Salmon) spore mass and, in some sori with 
an otherwise pale spore mass, spores towards the base of the sorus are this darker 
colour. The spore mass is at first agglutinated but later powdery. There is a 
central columella of host vascular tissue. 


Spores are mainly globose to subglobose, some oval, and a few slightly 
more angular in shape. Some tend to adhere in clumps of 10-15 or more, 
especially in the lighter coloured spore masses, and spores from such masses 
are usually more angular in outline than single spores. Spores from pale spore 
masses also have a paler golden yellow to greenish yellow (Munsell 10Y 9/6-8, 
2-5GY 9/4; Ridgeway Pale Green-Yellow) wall than those from the darker 
spore masses, where the spore wall is darker yellow to light golden brown (Munsell 
5Y 9/8, 7-5Y 9/10; Ridgeway Buff Yellow to Light Orange Yellow). Spores 
measure (15) 16-22 (25) um in diameter, with an occasional giant spore up to 
27 um. Oval spores are 17—23 (25) x12-17 um, and make up less than 10° 
of the number of spores. 


In some sori, fragments of hyaline hyphae 1-5-4 ym wide are seen amongst 
the spores. Often they are branched and twisted, and they vary in width along 
their length. Several spores with hyphal fragments up to 10-12 »m long attached 
to them occur and some spores have two fragments attached on opposite sides. 
The hyphae in the sori and attached to spores are seen mainly in paler, more 
agglutinated, less mature sori. 


The spore wallis rough. In equatorial view (Plate x, Fig. A) the roughening 
is seen as conical blunt projections, 1-5—2-0 um high and 1-5 um wide at the 
base. In surface view (Plate x, Fig. B; Text-fig. 1) these projections are seen 
to be the ridges of a reticulum, which is sometimes complete and produces angular 
areolae 1-5—-2-0 (2-5) um across. Often several ridges do not join up and an 
incomplete reticulum is formed. In a minority of spores, the surface pattern 
is seen aS elongated, sometimes branched ridges, which seems to represent a 
very incomplete reticulum (Text-fig. 1). These do not seem to be developmental 
stages as all patterns have been seen on lighter and darker spores in both 
agglutinated and powdery sori. 


Explanation of Text-fig. 1. 
Text-fig. 1. 


Figs. A-D. Surface markings of Ustilago cyperi-lucidi spores, showing complete and 
incomplete reticulate patterns. 


Figs. E-F. Details of reticulum. 
Figs. G-H. Spores germinating by septate promycelium producing sporidia. 


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J. WALKER 


logim 


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102 UNDESCRIBED SPECIES OF USTILAGO CYPERUS LUCIDUS R.Br. 


Germination occurs within 24 hours when dry spores are sown on potato 
dextrose agar plates at room temperature (fluctuating 15°-21°C. during the 
test). A four-celled promycelium from 26-33 ym long and 5-9 um wide is 
formed and each cell produces oval sporidia 7-10 x 4-6 wm (Plate x, Fig. EH; 
Text-fig. 1, G—H). 

Plants from the type collection have been grown on in the glasshouse. 
Basal shoots from infected parent plants gave rise to daughter plants with smutted 
florets, indicating that the smut systemically infects its host. 


RELATIONSHIP TO OTHER SPECIES 


There are comparatively few species of Ustilago with a light coloured 
(yellowish brown to light reddish brown) spore powder. The seed smuts of 
Cruciferae (Ustilago thlaspeos (Beck) Lagerh. in Syd., and others) and of Oxalis 
spp. (Ustilago oxalidis Ell. and Tracy), and a few species on hosts in the families 
Junecaceae and Cyperaceae fall into this group. Nannfeldt (1959) suspects that 
the crucifer smuts and U. oxalidis show some relationship to those species of 
Thecaphora which have a pale spore powder. The pale reddish-brown colour 
of the spore powder in some sori of U. cyperi-lucidi is somewhat lighter than 
that seen in spore powders of Thecaphora deformans Dur. and Mont. in Tul. 
(DAR 11070: spore mass reddish-brown, Munsell 10R 4-6/10) and 7. seminis- 
convolouli (Desm.) Liro (DAR 13830: spore mass reddish-brown, Munsell 
10R 4/8) but is very similar to that seen in a specimen of U. oralidis (DAR 11080 : 
pale reddish-brown, Munsell 2-5YR 6/12). In U. cyperi-lucidi, there is no sign 
of permanent spore balls or of spores ornamented only on one surface, or of other 
characteristics currently held for the genus Thecaphora. Spores of U. cyperi- 
lucidi are larger than those of U. owalidis and U. thlaspeos and differ in 
ornamentation. 


On the family Cyperaceae, Fischer and Holton (1957) list 111 species of 
smut fungi in 16 genera, and many other names occur in the literature. In 
Fischer and Holton’s list there are only six species of Ustilago and one variety 
on hosts in this family and only one of these, Ustilago subnitens Schroter and 
P. Hennings, in Hennings (1896) resembles U. cyperi-lucidi. U. subnitens 
was described originally from Brazil on Scleria sp. near S. pratensis Lind., and 
has also been reported on S. hispidula Hochst. from Ethiopia by Zundel (1953). 
It was described as an ovary smut, and in his commentary Hennings (1896) 
states that only some florets in an inflorescence were infected. The sori remained 
long closed and were pale cinnamon in colour. The spores were described as 
subglobose to ellipsoidal, 12-18 x10-15 um, with a smooth yellowish-brown 
epispore. He stated that thin threads were sometimes present between the 
spores and attached to some of them. From the description, U. subnitens 
seems similar to U. cyperi-lucidi, differing in the smaller smooth spores. Unfor- 
tunately the type specimen is unavailable in Herb. B and was possibly lost during 
World War II (Dr. H. Scholz, personal communication). No specimens under 
the name U. subnitens were found in the National Fungus Collections at 
Beltsville, where Zundel’s collections are kept (Dr. C. R. Benjamin, personal 
communication). Thus, no direct comparison of U. subnitens and U. cyperi- 
lucidi can be made. U. cyperi-lucidi does not resemble any other smut described 
on Cyperaceae. 


On the family Juncaceae, there are three species of Ustilago with a yellowish- 
brown spore powder. These are U. capensis Rees on Juncus spp. in South 
Africa (Zundel, 1953), U. vwyckii Oudemans and Beijerinck in Oudemans (1895) 
on Luzula spp. in Europe and America (Fischer, 1953; Zundel, 1953) and 
U. abstrusa Malencon on Juncus gerardii Loisl. in France (Zundel, 1953). All 
have a yellowish to pale reddish brown spore powder, with reticulate spores in 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Waues, Vou. 96, Part 2 


J. WALKER 103 


the range 14—24 um in diameter. From the descriptions, they seem very similar 
to one another and to U. cyperi-lucidt. 

Hight specimens of U. vuyckii, including the type collection, have been 
studied. The spore mass develops in the capsules of Luzula spp. and surrounds 
three small structures which may be ovule remnants. It is pale yellowish-brown 
to reddish-brown (Munsell 2-5YR 7/10 to Munsell 7-5YR 8/6) and is very similar 
to the darker shade seen in some sori of U. cyperi-lucidi. The spore mass in 
some sori is firmly agglutinated, but in others is granular to powdery. Spores 
are roughly spherical to globose, pale yellow in colour, (14) 16-20 (22) ym in 
diameter. They are strongly and regularly reticulate with roughly polygonal to 
hexagonal areolae 2-3 (3-5) um across. In equatorial view, the reticulum is 
seen to project 2-3 um above the spore surface (Plate x, Figs. C-D). Hyphal 
fragments are present in the sori, and many spores with one or two short hyaline 
hyphal fragments attached to them were seen. Some spores with a complete or 
partial, very thin (1 wm) hyaline sheath were seen, and this may be related to 
the method of spore development described for this species (Liro, 1924 ; Seyfert, 
1927). Some immature spores inside hyphae were common in one specimen 
(Syd. Mye. germ. 2878). Spores were described by Oudemans (1895) as 
germinating by a simple or branched promycelium, 24 x 2-3 um, which produced 
colourless elliptical to ovate sporidia 3-5x2-5 um. 


U. cyperi-lucidi is very similar to U. vuyckti in several respects. It differs 
mainly in its spore ornamentation, its smaller sporidia and its host family. 

Specimens of U. capensis and U. abstrusa have not been seen. As described 
by Oudemans (1895) and Zundel (1938, as Cintractia ; 1953), U. capensis occurs 
on Juncus spp. in South Africa, and produces golden yellow powdery sori in the 
ovaries. Spores are globose, 15-16 um diam., with a three-layered reticulate 
wall. Little else is known about it. U. abstrusa occurs on Juncus gerardii 
Lois. in France and produces powdery sori in the ovaries. Spores are yellowish, 
16—22 x 15-20 um, and reticulate, with areolae 2-2-5 um across (Viennot-Bourgin, 
1956, as Cintractia). Both seem to be closely related to U. vuycku, which attacks 
plants in the same family. 

The Ustilago spp. on Juncaceae discussed above were all transferred to the 
genus Cintractia by Ciferri (1931, pp. 56, 72) because of the presence in their sori 
of a columella. They do not fit into this genus as it is currently defined and are 
best left in Ustilago. 

No smuts with light-coloured spores have been recorded previously in 
Australia on the families Cyperaceae or Juncaceae. Examination of the 
Australian collections of Luzula spp. and Scleria spp. in Herb. N.S.W. did not 
reveal the presence of any smuts. Brittlebank’s (1937-1940) record of Farysia 
olivacea (DC.) H. and P. Syd. (as Ustilago) on Cyperus lucidus is based on a 
Specimen of the smut on a species of Carex where the host was identified 
incorrectly (McAlpine, 1910, p. 158). 


THE ORIGINAL COLLECTION OF Cyperus lucidus 

The occurrence of a smut in several specimens identified as Cyperus lucidus 
prompted an examination of the original collection of this species. Some 
variation in the shape of the nut in C. lucidus has been suggested by some workers. 
In his monograph, Kiikenthal (1935) describes the nut of the typical variety of 
C. lucidus as “‘ obovata trigona obtusa ”’ and lists only the original collection 
made by Robert Brown, No. 5884. The majority of collections listed from 
Queensland, New South Wales, Victoria and Tasmania he places as C. lucidus 
var. sanguineo-fuscus (Nees) Kiikenth. and describes its nuts as ‘‘lineari-oblonga”’. 
Willis (1962), in listing C. lucidus for Victoria, states ‘‘ The most widely distributed 
variety, and only one represented in Victoria, has more remote glumes and 


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104 UNDESCRIBED SPECIES OF USTILAGO CYPERUS LUCIDUS R.Br. 


comparatively narrower nuts than in the typical Port Jackson form; it is 
distinguished as var. sanguineo-fuscus (Nees, ut sp.) Kukenth.”. Hooker (1858) 
describes the nut of C. sanguineo-fuscus Nees as ‘‘ sharply trigonous, linear- 
obovate, acute, white, smooth ”’ and lists two specimens. A duplicate of one of 
these, Gunn 956, in Herb. NSW has been examined and has healthy nuts as 
described by Hooker. At present, all specimens in Herb. NSW are filed under 
the name C. lucidus. The possibility that the wider nuts described for typical 
C. lucidus by some authors were in fact smutted was considered and an examina- 
tion of Robert Brown’s original collection was made. 


Several florets from the type sheet in the British Museum (Natural History), 
and a duplicate sheet from the same collection in the Herbarium, Royal Botanic 
Gardens, Kew, were examined. On both sheets, two inflorescences were present. 
The British Museum specimen showed no nuts in one inflorescence and normal 
trigonous nuts in the other; the Kew specimen showed no nuts in one inflor- 
escence, and smutted nuts in all florets of the other (Plate x, Figs. B, D). 
Robert Brown thus unknowingly collected this smut, and his original collection of 
C. lucidus, made in the Sydney-Newcastle area of N.S.W., consisted of a mixture 
of healthy and diseased inflorescences. The portion in the British Museum is 
healthy and can be used as lectotype in accordance with the recommendation 
made by Stearn (1960) for the selection of lectotypes of Brown’s species. In 
nut shape, there is no difference between the BM specimen of C. lucidus and 
normal specimens of C. lucidus in Herb. NSW. Duplicates filed in other 
herbaria should be checked to see whether they are normal or smutted. The Kew 
specimen of Ustilago cyperi-lucidi in Robert Brown’s gathering of C. lucidus 
is one of the earliest specimens of a plant-parasitic fungus from Australia. 


SPECIMENS EXAMINED. 


Ustilago cyperi-lucidi J. Walker—all on Cyperus lucidus R.Br.—QUEENSLAND : 
Buderim Mountain, C. T. White, April 1916, NSW 65047 (portion as 
DAR 19558). 


NeEw SoutH WALES: Narrabri West, J. L. Boorman, June 1907, NSW 22730 
(portion as DAR 16728); Walcha Road, Northern Tablelands, J. L. 
Boorman, Dec. 1912, NSW 65071 (portion as DAR 16731); in swamp, 
Brunswick Heads, B. G. Briggs 757, 25.v.1954, SYD without number 
(portion as DAR 17585); Upper Cudgegong River, HK. of Olinda, 15 m. 
E of Rylstone, H. S. McKee, 10.1.1953, NSW 65074 (portion as DAR 16730) ; 
duplicate of same collection, SYD without number (portion as DAR 17584) ; 
Port Jackson, Robert Brown 5884, 1802-1805, Herb. K without number 
(sori and slides as DAR 17581) ; Five Crossings, Australian Capital Territory, 
N. T. Burbidge (Herb. Australiense 6579), 1.iv.1960, NSW 65079 (portion 
as DAR 16729) ; about 2500 ft. level, Clyde Mtn., R. Carolin B145, 21.iii1.1957, 
SYD without number (portion as DAR 17586); property of R. Skellern, 
Wyndham, W. of Pambula, J. Hindle, 4.vi.1969, NSW 118232 (portion as 
DAR 17842); same locality, J. Hindle, 11.vii.1969, DAR 17587 (TYPE) ; 
China Wall, Tooma, E. J. McBarron, 4.1.1963, Herb. EH. J. McBarron 7330 
(portion as DAR 17731); same locality, EK. J. McBarron, 4.1.1963, Herb. 
E. J. McBarron 7333 (portion as DAR 17730). 


VIcTORIA: Little River, Cathedral Range, near Taggerty, H. T. Clifford, 
1950, MEL without number (portion as DAR 17583). 


Ustilago vuyckii Oud. and Beijr. in Oud.—NETHERLANDS: near Voorschoten, 
L. Vuyck, 22.v.1894, on Luzula campestris, Herb. L, TYPE, (slide as 
DAR 19759); Keukenhof, L. Vuyck, 31.v.1894, on LZ. campestris Herb. L 
(slide as DAR 19760). 


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J. WALKER 105 


GERMANY: Sydow, Mycotheca germanica 2878, Brandenburg, near 
Kahnsdorf, E. Fahrendorff, June 1933, on L. campestris, Herb. L 937.56—90 
(slide as DAR 19763); Sydow, Myc. germ. 3264, Brandenburg, Tiefensee 
near Werneuchen, H. Sydow, 10.v.1939, on L. pilosa, Herb. L 942.57-77 
(slide as DAR 19761) ; Sydow, Myce. germ. 3265, Brandenburg, Hangelsberg, 
K. Fahrendorff, 18.v.1939, on L. pilosa, Herb L 942.57-30 (slide as DAR 
19762); Sydow, Ustilagineen 311, Marienspring, P. Sydow, 6.vi.1904, on 
L. pilosa, Herb. L 910.253-449 (slide as DAR 19765). 

SWEDEN: Fijellnas in Herjedalia, G. V. Lagerheim, July 1897, on L. pilosa, 
Herb. L (slide as DAR 19766); duplicate of last collection, Sydow 
Ustilagineen 217, Herb. L 910.253-438 (slide as DAR 19764). 


Ustilago oxalidis Ell. and Tracy—UNITED STATES OF AMERICA: On Ovalis stricta. 
Park Lane, Virginia, W. H. Long, 1911, DAR 11080 (duplicate of BPI 3863), 


Thecaphora deformans Dur. and Mont. in Tul.—UNITED STATES OF AMERICA : 
on Desmodium nudiflorum, Virginia Polytechnic Institute, Montgomery Co., 
Virginia, A. B. Massey, 4.xi.1941, DAR 11070 (duplicate of BPI 71541). 


Thecaphora seminis-convolvult (Desm.) Liro—RUMANIA : on Convolvulus arvensis 
L., near Odorhei, Transylvania, K. Vanky Ust. 3, 8.ix.1956, DAR 13830. 


Cyperus lucidus R. Br.—NEW SoutH WALES: Port Jackson, Robert Brown 5884, 
1802-1805, Herb. BM (lectotype). The collections of Cyperus lucidus in 
Herb. NSW were examined but the healthy collections are not listed in 
detail. 


The Australian collections of Scleria spp. and Luzula spp. in Herb. 
NSW were also examined. 


DISCUSSION 


The genus Ustilago is the largest single genus of the Ustilaginales, with about 
one-quarter of the described species in it (Fischer and Holton, 1957). In 
proportion, the genus is poorly represented on the host family Cyperaceae. 
Few smuts have been recorded on Australian Cyperaceae, and Ustilago cyperi- 
lucidi is the first species of Ustilago recorded on the family in Australia. It 
occurs in eastern Australia from southern Queensland to Victoria, and may occur 
over a wider range. Its detection in the field would be difficult, and all collections 
of it made up to the present have been detected initially only by a detailed 
examination of inflorescences in botanical specimens. 


Ustilago cyperi-lucidi causes a subtle distortion of infected plants. Nuts 
are altered in shape and this affects the appearance of the florets. It is possible 
that the alteration of the shape of the nuts by the smut in some collections of 
C. lucidus has been partly responsible in the past for varieties with narrow and 
wide nuts being distinguished. Systemic infection could result in the persistence 
of diseased plants in a locality and in these individuals retaining their character- 
istic appearance. A similar infection by a smut (DAR 6971 and others) has 
been found in N.S.W. on species in the Restio gracilis complex and on R. dimorphus 
Rk. Br. (Johnson and Evans, 1963). In this case, characters of the inflorescence 
were altered and, in the past, manuscript names, as varieties, had been given to 
some diseased specimens. The occurrence of U. cyperi-lucidi in portion of Robert 
Brown’s original gathering of C. lucidus also points to the potential hazard of 
inconspicuous diseases in botanical taxonomy. 


From the description, the smut of Scleria spp. described from Brazil as 
Ustilago subnitens is the species which most resembles U. cyperi-lucidi. No 
specimens of U. subnitens were available for study, and its exact relationship 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Wates, Vou. 96, Part 2 


106 UNDESCRIBED SPECIES OF USTILAGO CYPERUS LUCIDUS R.Br. 


to the present species cannot be determined. The pale-spored smuts of Juncaceae 
are also very similar to U. cyperi-lucidt, differing in details of spore ornamentation 
and host family. 

The method of spore development in U. cyperi-lucidi has not been studied, 
but the attachment of hyphal fragments to the spores and their presence in the 
sori is very similar to that seen in several specimens of U. vuyckit. Spore 
formation in chains in hyphae and the development of clamp connections have 
been reported for U. vuyckw (Liro, 1924; Seyfert, 1927) and a study of spore 
development in U. cyperi-lucidi would be of value in establishing its relationship 
to other species. 


NOTE ON THE SPELLING ‘‘ VUYCKII ”’ 


Most authors who have mentioned Ustilago vuyckii have used the spelling 
‘““ vuyjeku”’ for the specific epithet. This spelling also occurs on the labels of 
some exsiccati of this species (Sydow, Mycotheca germanica 3264 and 3265). 
The spellings “ vuijcku” and “ vuyckiw”’ both occur in Oudemans’ (1895) 
original paper, but on the type packet it is spelt ‘‘ vwyckii”’. As the fungus was 
named after its collector, L. Vuyck, the correct spelling is ‘‘ vwyckit”. IT am 
grateful to Dr. R. A. Maas Geesteranus for confirming this opinion. The 
confusion in spelling may have arisen from the difficulty with some handwriting 
in distinguishing between ‘“‘y”’ and ‘ij ”’. 


ACKNOWLEDGEMENTS 


J am indebted to Mr. O. D. Evans, National Herbarium of New South Wales, 
for bringing this smut to my notice and for comments on the taxonomy of 
Cyperus spp.; to Mr. J. Hindle, District Agronomist, Bega, for collecting living 
infected plants; to Mr. E. McBarron, Veterinary Research Station, Glenfield, 
for allowing me to examine specimens of Cyperus lucidus from his private 
herbarium ; to Mr. J. H. Willis, National Herbarium of Victoria, for lending 
specimens of C. lucidus; to Mr. R. Ross, British Museum (Natural History), 
for florets from and a photograph of the lectotype of C. lucidus ; to the Director, 
the Herbarium, Royal Botanic Gardens, Kew, for lending a duplicate of Robert 
Brown’s original collection of C. lucidus; to Dr. R. A. Maas Geesteranus, 
Rijksherbarium, Leiden, for lending collections of Ustilago vuyckii; to Dr. H. 
Scholz, Botanischer Garten und Museum, Berlin-Dahlem, and Dr. ©. R. Benjamin, 
National Fungus Collections, Beltsville, for searching for specimens of Ustilago 
subnitens ; and to Mr. A. Searle, Biological and Chemical Research Institute, 
Rydalmere, for taking the photographs. 


References 


BRITTLEBANK, C. C., 1937-1940.—A List of Australian Fungi. 410+5 pp. A manuscript list 
issued as blueprint copies by C.S.I.R.O. 

CrrERRI, R., 1931.—Quinta contribuzione allo studio degli Ustilaginales. Ann. Myc., 29: 1-74. 

FiscHer, G. W., 1953.—Manual of the North American Smut Fungi. xiit+343 pp., 136 figs. 
Ronald Press Co., New York. 

FiscHer, G. W., and Hotton, C. 8., 1957.—Biology and Control of the Smut Fungi. x-+-622 pp., 
107 figs. Ronald Press Co., New York. 

Hennines, P., 1896.—Beitrage zur Pilzflora Stidamerikas. I. Myxomycetes, Phycomycetes, 
Ustilagineae und Uredineae. Hedwigia, 35: 207-262. 

Hooker, J. D., 1858.—Flora Tasmaniae, Part 2, p. 80, Plate 139. 

Jounson, L. A. 8., and Evans, O. D., 1963.—A revision of the Restio gracilis complex. Contrib. 
N.S.W. Nat. Herb., 3 (4): 200-217. 

KUKENTHAL, G., 1935.—Cyperaceae-Scirpoideae-Cypereae. Das Pflanzenreich, IV, 20, 671 pp. 
(Reprint edition, 1956. J. Cramer, Stuttgart.) 

Lansouw, J., and Stariev, F. A., 1964.—Index Herbariorum. Part 1. The Herbaria of the 
World. 5th Edition. Regnum Vegetabile, 31: vi+251 pp. International Bureau for Plant 
Taxonomy and Nomenclature, Utrecht. 


PROCEEDINGS OF THE LINNEAN Socirry or New South WALES, Vou. 96, Part 2 


Proc. Linn. Soc. N.S.W., Vol. 96, Part 2 PLATE X 


it 


r es ae. a . “HP tat Pr 


ea 
: bi ie De ais 


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aihe Ro ee ’ 


wats Ryeeyie ’ iia} j my has ry W Gos a, an aes ‘ih hee 


Proc. Linn. Soc. N.S.W., Vol. 96, Part 2 PLATE Xi 


—_ 


fy iG 5 ia (i> | tal heel Ruan 


vi ie atte ‘ee 


yf ee ae m. ae, a F eee . 
# y a ba F | i Pitas, fae aie OMS fae a Cue oh a 
a ete Lee - » ‘ \ hyd y=), ‘ Boas cae : pr 
cS Ok ee a Bt pt ee ee MMOLE RGN 6 nal 
Konig y 4 a a 7 (ou Wy 3 
a a a, " 7 oo a — wees Vat } . _ Sie i a 


J. WALKER LOT 


Lrro, J. I., 1924.—Die Ustilagineen Fimnlands, I. Ann. Acad. Sci. Fenn., Ser. A, 17: 
xvii+636 pp. Helsinki. 

McAtpiIngE, D., 1910.—The Smuts of Australia. Their Structure, Life History, Treatment and 
Classification, vi--288 pp., 66 plates. J. Kemp, Government Printer, Melbourne. 

Munsell Book of Color, 1967.—-Neighbouring Hues Edition, Matte Finish Collection. Munsell 
Color Co., Baltimore. 

NANNFELDT, J. A., 1959.—Appendix, pp. 147-160, in Lindberg, Brita—Ustilaginales of Sweden 
(exclusive of the Cintractias on Caricoideae). Symb. Bot. Upsal., 16 (2): 1-175. 

Oupemans, C. A. J. A., 1895.—Over twee nog onbekende fungi: Septoria dictyotae en Ustilago 
vuycku. Verslagen van de Zittegen der Wis- en Natuurkundige Afd. van de Koninklijke 
Akademie van Wetenschaffen Amsterdam, 3: 54—57. 

Ripeeway, R., 1912.—Color Standards and Color Nomenclature. 1v--44 pp., 53 plates. 
Washington, D.C. 

SEYFERT, R., 1927.—Uber Schnallenbildung in Paarkernmyzel der Brandpilze. Zeitschr. f. 
Bot., 19: 577-601. 

STEARN, W. T., 1960.—An Introduction to Robert Brown's * Prodromus Florae Novae Hollandiae ~ 
pp. V—LII in the J. Cramer facsimile reprint, 1960. J. Cramer, Weinheim. 

VIENNOT-BouRGIN, G., 1956.—Mildious, oidiums, caries, charbons, rouilles des plantes de France. 
Texte et Atlas. HEncycl. Mycol., 26, 27. Paul Lechevalier, Paris. 

Wits, J. H., 1962.—A Handbook to Plants in Victoria. Vol. 1, Ferns. Conifers and Mono- 
cotyledons. xv+448 pp. Melbourne University Press. 

ZUNDEL, G. L., 1938.—The Ustilaginales of South Africa. Bothalia, 3 (3): 283-330. 

ZUNDEL, G. L., 1953.—The Ustilaginales of the World. xi+-410 pp. Contrib. No. 176, Dept. 
Botany, School of Agriculture, Pennsylvania State College. 


EXPLANATION OF PLATES 
PLATE Ix 
Ustilago cyperi-lucidi on Cyperus lucidus. A, left, healthy florets ex NSW 77363; right, 
smutted florets ex NSW 22730, x8. B, smutted florets from Robert Brown 5884 ex K, x8. 
C, left, three normal nuts ex NSW 77363; right, three smutted nuts ex NSW 22730, x13. 
D, left, three normal nuts from Robert Brown 5884, ex BM : right, three smutted nuts from Robert 
Brown 5884 ex K, one cut to show spore mass, x 13. 


PLATE X 
A-B, Ustilago cyperi-lucidt, equatorial and surface views of spores showing fine reticulum 
and irregular markings ex DAR 17587a, x1,000. C-—D, Ustilago vuyckii, equatorial and surface 
views of spores, showing coarser regular reticulum, ex TYPE in Herb. L, x1,000, &, Ustilago 
cyperi-lucidi, germinating spores with septate promycelium and sporidia, ex DAR 17587a, x 400. 


PROCEEDINGS oF THE LINNEAN Society oF NEw SourH Waters. Von. 96, Part 2 


SHOOT BLIGHT OF EUCALYPTUS SPP. CAUSED BY AN 
UNDESCRIBED SPECIES OF RAMULARIA 


J. WALKER AND A. lL. BERTUS 


Biology Branch, Biological and Chemical Research Institute, 
N.S.W. Department of Agriculture, Rydalmere 


(Plates XII, XIII) 


[Accepted for publication 17th February 1971] 


Synopsis 

Ramularia pitereka n. sp. which causes a shoot blight and leaf spot of young plants of Hucalyptus 
spp. is described. It has been found only in eastern New South Wales on plants growing in 
nurseries and glasshouses. Inoculation tests and observations in nurseries indicate that only 
Eucalyptus spp. in the series Corymbosae-Peltatae of Blakely are susceptible. Angophora costata 
seedlings have been artificially infected. The possibility that this may have some taxonomic 
significance in the two host genera is mentioned. A possible relationship between R. pitereka 
and the canker fungus of Hucalyptus ficifolia in Western Australia is discussed. 


INTRODUCTION 


A sheot blight of Hucalyptus seedlings has been known in New South Wales 
for at least fifteen years. The only published information on it reports an 
outbreak in the N.S.W. Forestry Commission’s nursery at West Pennant Hills 
in the autumn, winter and spring of 1960, and states: ‘‘ An unidentified fungus 
was found causing severe blighting of the young shoot of seedlings of L. maculata 
and over 50°% of some plantings of this species had been lost through the disease. 
The same fungus was found on a few seedlings of EL. eximia”’ (N.S.W. Department 
of Agriculture, 1961). The disease was almost certainly described earlier than 
this by Mrs. J. de Bavay (personal communication) in notes she prepared in 1955. 
She observed a shoot blight on Eucalyptus maculata Hook. seedlings growing in 
a Sydney glasshouse and described it as follows: ‘‘ Distortion was considerable 
in the young leaf and shoot tissue, in the infected zones. Scattered irregular 
necrotic areas appeared, over which a greyish white waxy bloom developed, due 
to the converging of masses of individual minute erumpent conidial pustules, 
each about 4 to } mm. in diameter.’’ This is very similar to the disease described 
below. 


Forestry officers have noticed the disease for several years on young plants 
in nurseries. It has been seen only on plants over three months old and, if not 
controlled by spraying, can kill or severely damage them. Although more 
common in spring and autumn, it has been collected throughout the year in 
nurseries, but has not been found on plants in the field. 

Recent work has shown that the causal fungus is'a previously undescribed 
species of Ramularia, which is described below. Standard abbreviations as 


given by Lanjouw and Stafleu (1964) are used in citing specimens in various 
herbaria. 


DESCRIPTION 
Ramularia pitereka sp. nov. (etym., pitereka, alba, in lingua aboriginum Australiae). 


Pustulae in surculis juvenilibus tortis albae, confertae, usque ad 100 ym 
diametro, in caulibus erumpentes, in foliis praecipue hypophyllis et ex stomatibus 
orientes. Conidiophora in strato denso, cum stromate basali hyalino ex hyphis 
intertextis constata ; stromata 15-50 ym crassa. Conidiophora hyalina, usque 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wass, Vou. 96, Part 2 


J. WALKER AND A. L. BERTUS 109 


ad 50 um longa, 2-2-5 um lata, plerumque cicatricibus numerosis parvis praedita, 
saepe geniculata, in regione cicatricum non septata. Conidia acrogena, levia, 
non septata, formae valde variabilis, plerumque clavata vel elongato-clavata, saepe 
cylindrica vel anguste pyriformia, conidia minoria saepe obclavata vel ovalia, 
(5) 6-17 (20) x2-5-5 (6-5) wm, in catenis brevibus interdum portata, cicatrice 
vel basali vel in quoque extremo praedita, per hypham simplicem vel per 


ABSA S AIS Rene saese Itura 
entro 
sima, 

CORRIGENDA 0 um 
eS ieee spite 

P. 109.—For Plate XI, read Plate XII. ae: 
For Plate XII, read Plate XIII. eae? 
P. 110.—For Plate XII, read Plate XIII. ot E. 

P. 111.—For Plate XII, read Plate XIII. 

P. 115.—For Plate XI, read Plate XII. on 3 
For Plate XII, read Plate XIII. ; and 
white 
Waxy 


cuticle and eventually rupturing it. The erumpent white pustules are up to 
100 um in diameter and closely packed on the diseased area (Plate x1, Fig. B). 
Some fusion between neighbouring pustules may occur during their development. 
On leaves, spots range from 1mm. to 2mm. in diameter up to large irregular 
areas which often develop along one edge and result in twisting and distortion 
of the leaf (Plate x1, Figs. C-E). Large infected areas often develop along the 
mid-vein (Plate x1, Fig. C). Spots are brown with a thin reddish-purple margin. 
Sunken brown lesions up to 1-5 em. long have been seen on petioles and stems, 
especially on Hucalyptus ficifolia F.Muell. (Plate x1, Fig. F). Sporulation 
occurs abundantly on all diseased tissues; on leaf spots it is more prominent 
on the abaxial surface. 


Pustules are composed of a dense layer of conidiophores borne on a 
plectenchymatous stroma up to 50 um thick, which develops in the host tissue 
(Plate xm, Fig. B). On shoots, the pustules rupture the epidermis and the thick 
cuticle, and are separated by their fragments ; on leaves, they are mainly hypo- 
phyllous and the conidiophores emerge in clusters through the stomates from a 
sub-stomal stromata. Conidiophores are hyaline, up to 50 ym long and 2-2-5 ym 
wide, and non-septate. Older conidiophores have many spore scars (up to 14 
or more), especially along the upper half, and are often prominently geniculate 
(Plate x11, Fig. E; Text-fig. 1, A and B). 

Conidia are borne singly at the tip of the conidiophore which grows on 
around the scar. The distance between scars is often very short, so that several 
conidia may appear clustered on a short length of conidiophore. Conidia are 
hyaline, smooth, strongly vacuolate, very variable in size and shape, generally 
clavate to elongated-clavate, often cylindrical to narrowly pyriform, smaller 
ones obclavate to oval, and measure (5) 6-17 (20) x 2-5-5( 6-5) um (Plate xu, 
Figs. C-E; Text-fig. 1). Whilst conidia on all hosts are variable in size and 
shape, there is a tendency for slight variations between hosts to occur. For 
example, many conidia from H. maculata are slightly longer and more regularly 
clavate than the somewhat shorter and more variable spores from E. eximia 
Schau. They often form short chains and have either a basal spore scar or scars 
at each end. Conidia sometimes germinate in the pustule, and they often form 
short secondary conidiophores bearing small secondary conidia (Text-fig. 1, A). 
Germination usually occurs by one to two germ tubes anywhere on the spore, 
except that germination through the scar has not been seen. 


PROCEEDINGS OF THE LINNEAN SocrETy oF NEw SoutH Wates, Von. 96, Part 2 


110 SHOOT BLIGHT OF EUCALYPTUS SPP. 


On agar media, colonies are white, raised in the centre and when older, often 
wrinkled by fine radial folds (Plate x1, Fig. A). They are covered with an 
abundant dry growth of conidiophores and conidia. Conidiophores are of 
indeterminate length. Spore-bearing areas up to 120 um long have been seen 


Jo G0U0" 
(boy 000, 


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Yext-fig. 1. A-—D, Ramularia pitereka conidia and conidiophores. A, from Hucalyptus 
eximia (DAR 19773), two conidia showing secondary conidiophores. B, from culture ex 
DAR 19773, showing types of conidiophore development, and three spores with vacuolate contents. 
C, from EH. maculata (DAR 19769), natural infection. D, from E. maculata (DAR 19770), artificial 
inoculation. EH, “ Sporotrichum destructor ”’ spores from IMI 7368. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 96, Part 2 


J. WALKER AND A. L. BERTUS atl 


and often several zones of scars occur along a conidiophore. Because of these 
zones of sporulation, conidia appear to be borne in clusters of varying size along 
the conidiophore. The arrangement of scars on the conidiophore is very similar 
to that described and illustrated by Hughes (1951, Fig. 2, E) for Acrotheca acuta 
Grove in culture. Often, conidiophores are branched and, apart from the spore 
scars, are very similar to vegetative hyphae. Conidia from all hosts are similar 
in culture and show a similar variability in shape to those from infected plants, 
but are generally slightly smaller, measuring (4) 5-13 (16) x (2-5) 3-5 (6) um. 
Short chains are often formed (Plate xm, Fig. C; Text-fig. 1). 


ISOLATION AND GROWTH IN CULTURE 


Ramularia pitereka is isolated easily from spores. It grows slowly on potato 
dextrose and malt agars and has an optimum temperature for radial growth 
between 20° and 25°C. After 24 days, colonies on malt agar are 12 mm. in 
diameter at 25° C. and 6 mm. in diameter at 30°C. No growth occurs at 35° C. 
and growth is very slight at 5° C., colonies being only 2 mm. in diameter after 
45 days. Colonies are white above and pale cream below. In cultures three to 
four months old, colonies often develop a faint cream to pink tint, and the 
medium is slightly darkened. 


INFECTION AND Host RANGE 


Artificial inoculations were carried out on a range of species of Hucalyptus 
and on Angophora costata (Gaertn.) Druce. Conidia from cultures were suspended 
in sterile water and applied with a fine brush to young shoots of seedlings in the 
glasshouse. Plants were kept in a humid atmosphere for 24 days after inoculation 
and then placed on the glasshouse bench. The characteristic symptoms of the 
disease and sporulation of the fungus were seen usually within three to four weeks 
of inoculation. The range of species tested and the results obtained are shown 
in Table 1. 


TABLE | 
Reaction of Seedlings of Eucalyptus and Angophora to Inoculation with Ramularia pitereka 


Section* Series Species Number Number 
Inoculated Infected 


Macrantherae Corymbosae-Peltatae  H. exvmia Schau. 10 2 
E. ficifolia F. Muell. 10 10 
E. maculata Hook. 10 10 
Transversae EH. longifoka Link and 10 0 
Otto 
E. punctata DC. 10 0 
Exsertae E. camaldulensis Dehn. 10 0 
Macrantherae (Normales) Globulares E. bridgesiana R. T. 10 0 
Bak. 
E. pulverulenta Sims 10 0 
Renantherae (Normales) Pseudo-stringybarks EH. pilularis Sm. 10 0 
Pachyphloiae E. caliginosa Blakely 10 0 
and McKie 
Porantheroideae (Nor- Buxeales E. albens Benth. 10 0 
males) 
Terminales _— ' E. sideroxylon A. Cunn. 10 0 
ex Woolls 
Platyantherae Subulatae E. flocktoniae Maiden 10 0 
Angophora costata 20 15 


(Gaertn.) Druce 


* The Hucalyptus spp. are arranged according to the sections and series given by Blakely 
(1965) and their nomenclature follows Johnston and Marryatt (1965). 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wa tes. Vou. 96, Part 2 


112 SHOOT BLIGHT OF EUCALYPTUS SPP. 


In these tests the only Hucalyptus spp. attacked were in Blakely’s (1965) 
series Corymbosae-Peltatae. Several plants of Angophora costata were also 
infected. The significance of these results is discussed below. 


POSSIBLE RELATIONSHIP TO CANKER FUNGUS OF FUCALYPTUS FICIFOLIA 


Possibly closely related to R. pitereka is the fungus that causes canker 
disease of adult trees of red-flowering gum (H#. ficifolia F. Muell.) in Western 
Australia. Little has been published about this organism. In 1936 Mr. H. A. 
Pittman, then Government Plant Pathologist, reported on a canker disease of 
EL. ficifolia in Western Australia (Anon., 1936). It had apparently been known 
for some years before this (Beard, 1963 ; Cass Smith, 1970), but Pittman was the 
first to isolate the causal fungus and suggest control measures. At the time he 
also sent a culture to the Commonwealth Mycological Institute. Subsequently, 
the fungus was referred to in various publications as ‘‘ Sporotrichum destructor 
Pittman ”’. MacNish (1963) lists it as ‘“‘ Sporotrichum destructor Pittman n. sp.” 
and in a footnote states ‘“‘ pathogen named 1936”. Later authors dealing with 
canker have used this name (Beard, 1963; Cass Smith, 1970) or have referred 
to the disease as Sporotrichum canker (Pryor and Willing, 1963). 


The name ‘“ Sporotrichum destructor’’ has never been validly published, 
and no description of the fungus has been found in the literature. Mr. Pittman 
(personal communication) wrote that, whilst he proved its pathogenicity and 
tentatively referred to it as Sporotrichum, he did not eall it ‘ S. destructor”. 
However, a description under this name has been obtained from Mr. H. L. Harvey, 
of the Department of Agriculture, Perth, Western Australia. This description, 
dated 20.x.1936, came from files of correspondence with the Commonwealth 
Mycological Institute, and was prepared there by Mr. EK. W. Mason (F. C. 
Deighton, personal communication). It describes the fungus in culture. Colonies 
are described as ‘‘ floccose, then powdery, finally showing radiating striations ”’, 
at first white and later pinkish-buff in colour. The fertile hyphae are described 
as ‘‘ usually long (up to 150 u), more rarely lateral and shorter, unbranched, 
cylindrical, typically straight and bearing in a stellate manner a single whorl, 
or occasionally a spiral of a few (up to 10) conidia, sometimes gently bent and 
provided with two or even more verticills or spirals. Conidia hyaline, cylindrical 
or oblong, rounded at both ends, 0-septate, sessile (that is—without sterigmata), 
6-8 uw long, 2 to 2-5 » broad.” 

The only specimen filed under the name “* Sporotrichum destructor’ at the 
Commonwealth Mycological Institute has been examined. It is a dried malt 
agar culture, light buff (Ridgeway, 1912) in colour, now showing as a fairly 
compact felted mat of hyphae on the dried agar. Abundant hyaline, unicellular 
conidia are present. They are variable in shape, cylindrical to clavate, with 
smaller conidia oval to obovate, measuring (2-5) 3-10 (12-5) x (2) 2-5-3-5 (4) wm, 
and have an indistinct spore scar at one end. Evidence of chain formation was 
present aS some spores had a sear at each end. Conidiophores could not be 
clearly distinguished, but some fragments with probable indistinct spore scars 
were seen. In some cases, clusters of conidia were seen around probable 
conidiophores. 

The spores from this specimen are smaller than those of R. pitereka in culture. 
From Mr. Mason’s description, its cultural characters and some aspects of its 
conidiophore and conidium morphology are similar to those of R. pitereka, but 
until it can be studied from fresh specimens, its identity remains uncertain. 


SPECIMENS EXAMINED 


Ramularia pitereka J. Walker and A. L. Bertus—NEw SouTH WALES: on 
Eucalyptus eximia Schau., in nursery, Forestry Commission of N.S.W., 
West Pennant Hills, A. L. Bertus and J. Walker, 3.vi.1970, DAR 19773 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Waters, Vou. 96, Part 2 


J. WALKER AND A. L. BERTUS 113 


(type) (portion filed as IMI 151369); same locality, J. Walker, 18.xi.1960, 
DAR 6469; same locality, A. L. Bertus, 5.v.1970, DAR 20366 ; artificial 
inoculation, glasshouse, Department of Agriculture, Rydalmere, A. L. 
Bertus, 15.xi.1967, DAR 19771. On Eucalyptus ficifolia F.Muell., in 
nursery, Forestry Commission of N.S.W., West Pennant Hills, A. L. Bertus 
and J. Walker, 3.vi.1970, DAR 20361. On Eucalyptus maculata Hook., 
in nursery, Forestry Commission of N.S.W., West Pennant Hills, J. Walker, 
30.v1.1960, DAR 5867a ; in commercial nursery, Beverley Hills, J. Stronach, 
24.11.1967, DAR 19768 ; same locality, J. Stronach, 28.11.1967, DAR 19769 
(portion filed as IMI 151368) ; artificial inoculation, glasshouse, Department 
of Agriculture, Rydalmere, A. L. Bertus, 15.xi.1967, DAR 19770; in 
commercial nursery, Glenhaven, A. L. Bertus, 5.xi.1970, DAR 20468. 
On Angophora costata (Gaertn.) Druce, artificial inoculation, glasshouse, 
Department of Agriculture, Rydalmere, J. Walker, 16.xi.1970, DAR 20469. 


‘* Sporotrichum destructor ’—-WESTERN AUSTRALIA: in dried malt agar culture, 
H. A. Pittman, 1936, IMI 7368 (slides filed as DAR 20410). On Hucalyptus 
ficifolia F. Muell., King’s Park, Perth, L. Harvey, Sept., 1970, DAR 20409 
(cankers only, no fungus seen). 


DISCUSSION 


No previous description of a species of Ramularia on hosts in the family 
Myrtaceae has been found. Most species of Ramularia occur on herbaceous 
hosts and generally cause comparatively minor damage such as leaf spots and 
blotches. Rk. pitereka occurs on young growth of woody hosts and causes more 
severe damage than is usually seen with many other Ramularia spp. Whilst 
comparatively few species of Ramularia have been studied in culture, those 
that have generally shown sparse sporulation on artificial media; however, 
hk. pitereka produces abundant spores in culture. 

In the limited inoculation tests carried out so far only Hucalyptus spp. 
in the series Corymbosae-Peltatae of Blakely (1965) have been susceptible. 
The species used have also been arranged (Table 2) according to a new classi- 
fication and coding of the genus (Pryor and Johnson, 1971) and in this system 
all the susceptible species fall into the subgenus © Corymbia. The one species 
of Angophora tested, A. costata, waS experimentally infected. It is possible 
that this may have some taxonomic significance in the two host genera, and tests 
on a wider range of species will be carried out. The affinity in essential oil and 
certain morphological characters between species of Hucalyptus in the above and 
related series and the genus Angophora has been pointed out previously (Baker 
and Smith, 1920). 

Finally, R. pitereka may be the same as the undescribed fungus causing 
canker of red-flowering gums in Western Australia, and referred to in the literature 
as ‘‘ Sporotrichum destructor Pittman’. If so, it may have been introduced 
into eastern Australia from the west. Cass Smith (1970) indicates that in 
Western Australia the disease occurs naturally on Eucalyptus calophylla R. Br. 
ex Lindl., and that infections on cultivated H. ficifolia probably originated from 
naturally infected E. calophylla. So far, no plants infected with R. pitereka 
have been found in naturein N.S.W. Cass Smith (1970) also lists EH. haematoxylon 
Maiden as another host of the canker fungus. All the known hosts of ‘‘ Sporo- 
trichum destructor” thus occur in Blakely’s series Corymbosae-Peltatae (or 
subgenus © Corymbia in Pryor and Johnson’s (1971) classification), and this 
provides a further indication of a possible relationship between the canker 
fungus and R. pitereka. 


PROCEEDINGS OF THE LINNEAN Soctety oF New SoutH Wates. Vou. 96, Part 2 


SHOOT BLIGHT OF EUCALYPTUS SPP. 


114 


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PROCEEDINGS OF 


Proc. Linn. Soc. N.S.W., Vol. 96, Part 2 PLATE XII 


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Proce tins, Soc. NISIWe, Vol: 96) Part 2 PLATE XIIt 


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HERB. 1.M.I. 
COMMONWEALTH MYCOLOGICAL INSTITUTE 


Name ..Sperotrichum destructor. Fittman oo! 
Host 290... OK. BBL Y PHS oo cccccsneseessneenseenneesenseee 
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Dried dowm cult. on Malt Agar- one of the sbenltrvres 


from which the diagnosis was drawn up, 
‘SY PUPP PRT EUDIT PUTED T TVA] TS] CTE CUTE TE CEPT EP) 


J. WALKER AND A. L. BERTUS 115 


ACKNOWLEDGEMENTS 


We are indebted to Mr. D. J. Hartigan, Forestry Commission of N.S.W., 
and to the officers at the Forestry Nursery, West Pennant Hills, for providing 
information on the disease and specimens. Mrs. J. de Bavay, of Armidale, 
N.S.W., made available her early observations and Mr. H. A. Pittman, Upper 
Hawthorn, Victoria, provided valuable information on his work with “ Sporo- 
trichum destructor’. We thank Mr. F. C. Deighton and Dr. M. B. Ellis, of the 
Commonwealth Mycological Institute, for comments on these fungi and lending 
the specimen of ‘“ S. destructor”. Mr. lL. Harvey, Officer-in-charge, Plant 
Pathology Branch, Department of Agriculture, Perth, sent us information on 
canker disease and provided a canker specimen, and Mr. L. A. S. Johnson, 
National Herbarium of N.S.W., sent seed of species of Hucalyptus and Angophora 
and information on their classification. Professor L. D. Pryor and Mr. Johnson 
allowed us to quote from their new classification and coding of the genus 
Hucalyptus. The photographs were taken by Mr. A. Searle, Department of 
Agriculture, Rydalmere. 


References 

ANon., 1936.—Red-flowered gums. Control of canker disease. Ex West Australian, 17 Sept., 
1936. (Abstract. in Rev. appl. Mycol., 16: 138, 1937.) 

Baker, R. T., and Smitu, H. G., 1920.—A Research on the Hucalypts Especially in Regard to Their 
Essential Oils. 2nd Edition. xv+471 pp. W. A. Gullick, Government Printer, Sydney. 

Brarp, J. S., 1963.—The stem canker disease of red-flowering gum. Australian Plants, 2 (14) : 
62-63. 

BuaKxety, W. F., 1965.—A Key to the Eucalypts, with Descriptions of 522 Species and 150 Varieties. 
3rd Edition. 359 pp. Forestry and Timber Bureau, Canberra. 

Cass SmituH, W. P., 1970.—Stem canker disease of red flowering gums. J. Dept. Agric. W. Aust., 
4th Ser., 11 (2): 33-39. 

Hucues, 8S. J., 1951.—Studies on micro-fungi. V. Acrotheca. Mycol. Pap. No. 38, 8 pp., 2 figs. 
Commonwealth Mycological Institute, Kew. 

JOHNSTON, R. D., and MarryatT, Rosemary, 1965.—Taxonomy and Nomenclature of Hucalypts. 
Leaflet No. 92. 24 pp. Forestry and Timber Bureau, Canberra. 

Langouw, J., and StarLeu, F. A., 1964.—Imdex Herbariorum. Part 1. The Herbaria of the 
World. 5th Edition. Regnum Vegetabile, 31: vi+251 pp. International Bureau for 
Piant Taxonomy and Nomenclature, Utrecht. 

MacNisu, G. C., 1963.—Diseases recorded on native plants, weeds, field and fibre crops in Western 
Australia. J. Agric. W. Aust., 4th Ser., 4 (6): 401-404, 407—408. 

N.S.W. DEPARTMENT OF AGRICULTURE, 1961.—Plant Disease Survey for the Year Ending 30 June, 
1961, p. 49. 

Pryor, L. D., and Jounson, L. A. 8S., 1971.—A classification of the eucalypts. 

Pryor, L. D., and Witting, R. R., 1963.—The red flowermg gum. Australian Plants, 2 (14) : 
35-37, 63-64. 

RipgEway, R., 1912.—Color Standards and Color Nomenclature. iv+44 pp., 53 plates. 
Washington, D.C. = 


EXPLANATION OF PLATES 
PLATE XI 
Ramularia pitereka on Hucalyptus spp. A, B, on H eximia (DAR 19773) showing shoot 
symptoms and white pustules. C, D, E, on H. maculata (DAR 5867a) showing shoot blight and 
leaf spotting and distortion. F, on EH. ficifolia (DAR 20361) showing stem lesion. A, C, D, E, 
slightly less than natural size. B, x3:5. F, x2. 


PLATE XII 
A-E, Ramularia pitereka. A, colonies 24 days old, on potato dextrose agar (natural size). 
B, young pustules on H. exwimia shoots, x 288. C, spores in culture, some forming short chains, 
<x 400. D, spores from EH. maculata (DAR 19769), «1,000. E, spores and fragment of conidio- 
phore from EH. eximia (DAR 19773), 1,000. F. G, “ Sporotrichum destructor’ (IMI 7368), 
spores (1,000) and specimen (scale in mm.). 


PROCEEDINGS OF THE LINNEAN Soctery or New South Wates, Vor. 96, Part 2 


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AUSTRALASIAN MEDICAL PUBLISHING CO. LTD. 
71-79 ARUNDEL ST., GLEBE, SYDNEY, N.S.W., 2037 


THE REACTION OF SEEDLINGS OF CITRUS SPP. AND RELATED 
GENERA TO PHYTOPHTHORA CITROPHTHORA 


P. BROADBENT, L. R. FRASER AND Y. WATERWORTH 
) 


Biological and Chemical Research Institute, Rydalmere, Department of 
Agriculture, N.S.W. 


[Accepted for publication 23rd June 1971] 


Synopsis 
Varieties and selections of Crtrus and related genera were inoculated with Phytophthora 
citrophthora to determine their resistance to root and collar rots. 


Poncirus trifoliata and its hybrids, Microcttrus australis and. Severinia buaifolia, were highly 
resistant to stem inoculations of P. citrophthora ; mandarins were intermediate in susceptibility, 
while Rangpur lime, sweet orange varieties, Smooth Sevilie and rough lemon were highly 
susceptible. Smooth Seville is moderately resistant to root rot. Ruby Blood 3118 and 
Parramatta orange 3443 were more resistant to Phytophthora root rot than other sweet orange 
selections tested. 


Of 150 clones of Poncirus trifoliata, all were uniformly and highly resistant to Phytophthora 
root rot. 


INTRODUCTION 

Phytophthora citrophthora (Sm. and Sm.) Leon, which causes root and 
collar rots, is one of the most destructive pathogens of citrus in New South 
Wales. The use of resistant rootstocks offers an excellent means of reducing 
the losses from root rot, and the resistant rootstock Poncirus trifoliata (L.) Raf. 
is widely used in New South Wales. However, it grows poorly in highly acid, 
highly alkaline or saline soils and is unsuitable for use as a rootstock for Hureka 
lemons (Citrus limon (l.) Burm.f.) because of its susceptibility to exocortis 
virus (scalybutt), which many old-line lemons carry. There is also an incom- 
patibility factor which causes a bud-union weakness (Nauriyal et al., 1958) with 
‘““Hureka’’ lemons. Therefore, an alternative Phytophthora-resistant stock 
without these disadvantages is needed and has been sought over a number of 
years. 

Field behaviour of citrus stock-scion combinations in root rot-liable soils 
is difficult to assess on the basis of laboratory, glasshouse or small field plot 
testing. Factors other than the intrinsic susceptibility of the rootstock to 
rotting of roots are involved. The reaction to Phytophthora spp. of the scion, 
the regenerative ability of the stock, the level and type of nutrition, and possibly 
other factors may modify this and the final assessment must be made on long- 
term field behaviour. However, initial screening can only be made on the basis 
of the amount of disease resulting from inoculation, and several methods have 
been employed to arrive at a satisfactory rating. 


Two species of Phytophthora are capable of causing root and collar rots of 
citrus, P. citrophthora and P. nicotianae (B. de Haan) var. parasitica (Dast.) 
Waterh. Their relative importance is dictated by temperature, P. citrophthora 
being the prevalent species at lower temperatures and P. nicotianae var. parasitica 
at higher temperatures. In New South Wales P. citrophthora is the predominant 
species. P. nicotianae var. parasitica is only rarely seen, and is restricted to the 
warmer parts of the central and north coast. 


The resistance of citrus species and varieties as reported from California, 
where both species of Phytophthora occur, and Florida and Queensland, where 


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120 REACTION OF SEEDLINGS OF CITRUS TO PHYTOPHTHORA CITROPHTHORA 


P. nicotianae var. parasitica is the predominant species, follows the same general 
pattern for both fungi, with only minor variation. No variation in pathogenicity 
of different isolates has been reported for P. citrophthora, but in Florida it has 
been found that considerable variation in pathogenicity occurs in P. nicotianae 
var. parasitica (Grimm and Whidden, 1962). 


Methods of testing for Phytophthora resistance vary. Klotz and Fawcett 
(1930), Fraser (1942) and Rossetti (1947) studied the resistance of a large number 
of species and varieties of Citrus and related genera to bark inoculation with 
P. citrophthora, and Klotz et al. (1958a) examined the resistance to P. nicotianae 
var. parasitica using bark inoculations. To assess the percentage decay of 
fibrous roots by P. citrophthora and P. nicotianae var. parasitica, Klotz et al. 
(1958b) immersed the root systems in aerated water cultures containing the 
fungus. Carpenter and Furr (1962) inoculated roots with P. nicotianae var. 
parasitica and then transferred the seedlings to incubation beds to determine 
their reaction to infection. Incorporation of inoculum in soil in which the test 
plants are established has been fairly widely used. The results of all these 
different approaches indicate that resistance occurs in Poncirus trifoliata and its 
hybrids and to a lesser extent in Citrus aurantium L. (sour oranges), and a range 
of susceptibility is shown by orange, mandarin, grapefruit, lemon and other 
clones. 


GENERAL METHODS 
Three methods of assessing resistance were used in this study : 
(a) Infestation of steam-air treated (140° F./30 min.) soil by adding corn- 
meal sand cultures of pathogenic isolates of P. citrophthora. 
(b) Growth of plants in Hoagland’s solution (Hoagland, 1919) to which 
cultures of P. citrophthora on lucerne (Medicago sativa L.) stems were 
added. 


(c) Inoculation of wounded stems. 


TABLE | 
Reaction of Citrus spp. and Related Genera to P. citrophthora in Soil 


Number of Plants with Collar Rot 


Common Name Botanical Name Mean Number 
Above Soil Below Root of 
Ground Level Ground Rot Plants 
Rating Kaulled 


Trifohate orange Poncirus trifoliata (L.) 0 0 0 0-8 0 
Raf. 
Rusk citrange P. trifoliata xC. sinensis 0 0 0 0-8 0 
Sampson tangelo C. paradist xC. reticulata 0 0 2 1-5 0 
Thornton tangelo C. paradisi x C. reticulata 1 0 I 1-5 0 
Harvard sweet orange C. sinensis (L.) Osbeck 1 1 3 Ilo) 2 
Cleopatra mandarin C. lycopersicaeformis Hort. 1 1 1 21 J 
ex Tan. 
Sathgudi orange C. sinensis (L.) Osbeck 0 2 2 2-1 il 
Emperor mandarin C. reticulata Blanco I 0 1 2-2 0 
Rough lemon (South C. jambhirt Lush. 2 4 2 2-2 2 
Africa) 
Kara mandarin C. reticulata Blanco 1 2 1 2-3 0 
Ellendale mandarin C. reticulata Blanco 1 0 1 2-3 0 
Rough lemon (N.S.W.) C. gambhiri Lush. 3 5) 3 2-5 I 
Volkamer lemon C. volkameriana Pasq. 4 5 8 2°5 1 
Ruby blood sweet C. sinensis (L.) Osbeck 1 1 2 27 1 
orange 
Karna Khatta C. karna Raf. 7 6 7 3-1 6 


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BROADBENT ET AL. IAS 


Citrus seeds were germinated in containers of steam-air treated soil 
(140° F./30 min.) When four to six inches tall, 10 seedlings of each variety were 
selected for uniformity and planted singly in one gallon containers, or in rows 
in large troughs of steam-air treated soil or transferred to aerated nutrient 
solution. 


HXPERIMENTAL 
(1) Reaction of Citrus spp. and Related Genera to Root Inoculations with P. 
citrophthora: In Soil 


Experiment 1: Citrus seedlings in one gallon containers were arranged randomly 
on the glasshouse bench. Ten replicates and 15 varieties were used. Ten gm. 
of inoculum was placed at a depth of 4 cm. in three holes around each plant. 
Inoculum consisted of a mixture of 12 isolates of P. citrophthora grown separately 
on corn-meal sand for six weeks. Hight of the 10 replicates were harvested 
after 11 weeks and the final two replicates 18 days later. 


The root rot rating of each plant was assessed on a scale ranging from 0 
(no observable injury) to 5 (most of root system rotted) and each rating represents 
the mean of 10 replicates. (Results are given in Table 1.) 


The hypocotyl region of plants of Karna Khatta, rough lemon, sweet orange 
and Volkamer lemon was particularly susceptible to infection by P. citrophthora 
(Table 1). Often severe collar infection occurred with little root damage. 
Mandarins and tangelos were less severely infected by P. citrophthora than 
sweet orange or lemon. In the two replicates examined at 14 weeks, growth of 
roots had recommenced after a winter dormancy, and the pioneer roots which 
had been produced were not attacked by P. citrophthora even though a very 
high concentration of inoculum was present in the soil. 


Experiment 2: The reaction to P. citrophthora of 21 sweet orange and 14 sour 
orange clones and a number of other .varieties was assessed. Seedlings were 
grown in one gallon containers and inoculated at 16 months with corn-meal sand 
culture of four isolates of P. citrophthora. Plants were examined after six months 
and a visual assessment made of root rot injury on a scale similar to that used in 
Experiment 1. (The results are given in Table 2.) 


In Aerated Nutrient Solution 


Experiment 1: Seedlings 6in. high were suspended in aerated Hoagland’s 
solution so that only the roots were immersed. A pathogenic isolate of 
P. citrophthora obtained from diseased citrus roots and grown on sterilized stems 
of iucerne, where it had produced sporangia, was enclosed in cheese-cloth and 
suspended in the culture solution for 24 hr. One month after inoculation the 
seedlings were removed from the Hoagland’s solution and placed in a 1°% solution 
of 2,3—-5-triphenyl tetrazolium chloride (Klotz and De Wolfe, 1965) for 24 hr. 
in the absence of light and assessed on the basis of stained as compared with 
unstained roots. 


Poncirus trifoliata, Troyer citrange, Ruby blood sweet orange, sweet orange 
(Symons clone), Smooth Seville (Warnock clone), Israel sour orange and rough 
lemon, in that order, showed increasing susceptibility to root rot as shown by 
stained living tissue. The method of testing susceptibility to Phytophthora 
root rot in aerated nutrient solution was a severe one, giving rapid results which 
were uniform for any given variety or species of citrus. 


(2) The Reaction of Citrus spp. and Related Genera to Stem Inoculations with 
P. citrophthora 


Seedlings, 18 months old, grown in one gallon cans, were arranged randomly 
on benches in the glasshouse in 10 replications. In Experiment 1, the resistance 


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122 REACTION OF SEEDLINGS OF CITRUS TO PHYTOPHTHORA CITROPHTHORA 


TABLE 2 
Reaction of Citrus spp., Varieties and Related Genera to P. citrophthora 


Mean 
Common Name Botanical Name Root Rot 
Rating 
Highly resistant : 
Trifohate orange 22* Poncirus trifoliata (L.) Raf. 1 
Resistant : 
Stuart Smooth Seville (10)+ C. aurantium L.? 1-5 
Roberts Smooth Seville (10) 55 .s ee, 1-6 
Appleby Smooth Seville PA a DENA? 1:6 
Haddon Smooth Seville an 45 Bey a 1:7 


Moderately susceptible : 


Parramatta orange 3443 CO. sinensis (lL) Osbeckx 2 

Ruby blood orange 3118 09 35 ees: 2-1 
Rough lemon, Terrigal No. 3 selection (10) ©. gambhirt Lush. Bo ih 
Rough lemon, Terngal No. 1 selection (10) i 53 an 2-2 
Warnock Smooth Seville C. aurantium Li. ? 22, 
Rough lemon, Terrigal No. 4 selection (10) ©. gambhiri Lush. 2-4 
Ellendale mandarm C. reticulata Blanco ? 29 
Citrus “ zigardia’”’ (a sweet orange type) Origin unknown 2°5 
Emperor mandarin (10) C. reticulata Blanco 2-6 
Marchant rough sour orange (10) C. aurantium L. 2°6 
Sweet orange 1259 C. sinensis (L.) Osbeck Qi] 
Haddon rough sour orange A (10) C. aurantium L. 2°7 
Haddon rough sour orange C (10) xf A 3M AO 7 
Haddon rough sour orange B (10) es A $5 2459) 
Rough lemon, Terrigal No. 2 selection (10) CC. gambhiri Lush. 2°7 
St. Michael orange 3109 (10) C. sinensis (L.) Osbeck 2-9 

Susceptible : 

Narara rough lemon C. jambhiri Lush. 3 

Cleopatra mandarin (N.S.W.) (10) C. lycopersicaeformis Hort. ex Tan. 3 

Brazilian sour orange C. aurantium L. aril 
Sweet lime (10) C. limettioides Tan. 3-2 
Wilson navel-type sour orange C. aurantium L. 3°3 
Chaffey sweet orange C. sinensis (L.) Osbeck 3-4 
Somersby rough sour orange A C. aurantium L. 3-5 
Somersby rough sour orange B a5 66 S 3-4 
Israel sour orange i Be Js 3:5 
Pineapple orange 3199 C. sinensis (L.) Osbeck 3-6 
Parramatta orange 3187 (10) es 6 ae Aes 3:6 
Bittersweet orange C. aurantium L. 3:7 
Parramatta orange 3188 C. sinensis (L.) Osbeck 3:8 
Homosassa orange 3104 as B x Rs 3:8 
Waikerie sweet orange 3185 fs be oe ae 3-9 

Highly susceptible : 

White Siletta orange C. sinensis (L.) Osbeck 4 

Yuzu (10) C. gunos Sub. ex Tan. 4 

Jaffa orange 3610 (10) C. sinensis (L.) Osbeck 4 

Mediterranean sweet orange sh %. sin i 4 

Cleopatra mandarin (ex Florida) C. lycopersicaeformis Hort. ex Tan. 4-1 
Campuda sweet orange 3613 (10) C. sinensis (L.) Osbeck 4-1 
Parramatta orange 3146 te BA a s 4-2 
Mayo orange ie ce ie Me 4-3 
Paterson orange 3449 oe 35 RC a5 4-4 
Calamondin (10) C. madurensis Loureire 4:6 
Maltese blood orange C. sinensis (L.) Osbeck 4-8 
Seedling sweet orange (Parramatta type) BS e, a ae 4-8 
Joppa orange AA & aa aA 4-8 


* Accession number referring to parent trees at the Department of Agriculture Horticultural 
Research Station, Narara. 


+ Number of plants tested was 20 unless otherwise stated. 


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BROADBENT ET AL. 123 


of seven varieties of mandarin, Ruby blood sweet orange and rough lemon were 
tested. In Experiment 2 two trifoliate orange clones, five trifoliate hybrids, 
nine sweet orange clones and nine miscellaneous citrus species were tested. 
Dises of bark (5 mm. diameter) were removed with a cork borer from the collar 
region of the stem 40mm. above soil level, inoculum inserted, and the area 
bound with plastic budding tape. Inoculum consisted of discs 5mm. diameter 
cut from a six-day-old agar culture of P. citrophthora newly isolated from diseased 
citrus roots. The wrappings were removed after three weeks in Experiment 1, 
and after 10 days in Experiment 2 and the lesion measured. One plant of 
each variety was treated similarly, but no inoculum was added. 


The reaction of mandarin varieties to collar rot is given in Table 3. Poncirus 
trifoliata and its hybrids, Microcitrus australis and Severinia buxifolia, were all 
highly resistant to stem inoculations of P. citrophthora (Table 4). The mandarins 
were intermediate in susceptibility, while Rangpur lime, sweet orange varieties, 
Smooth Seville and rough lemon were highly susceptible. There were no 
significant differences between sweet orange varieties (overall mean=2-03, 
S.E. (n'=76)= +0-08 on 67 degrees of freedom). Similarly, the two trifoliate 
selections did not differ significantly (overall mean =0-127,8.E. (nt'=11)= +0-0438 


TABLE 3 


Reaction of Citrus spp. and Related Genera to Stem Inoculation with 
Phytophthora citrophthora 


Length of 


Lesion 
Common Name Botanical Name in Cm. 
(Mean of 10 
replicates) 
Kara mandarin C. reticulata Blanco 0-92 
Searlet mandarin ss ae a 1-18 
Emperor mandarin =e fh 5 1-28 
Ellendale mandarin O. reticulata ? 1-36 
Dancy mandarin C. reticulata Blanco 1-49 
Rough lemon C. jambhiri Lush. Ihoisyss 
Acid mandarin C. reticulata Blanco 1-66 
North Coast sour mandarin aa 55 a 1-76 
Ruby blood sweet orange C. sinensis (L.) Osbeck 2-05 


Standard deviation: 0-451 em. or 30-:9%. 


Difference necessary for significance (59%) m comparison of 
means=0-451 cm. 


on nine degrees of freedom). One F, seedling of Yanco hybrid 3700 was highly 
susceptible to collar rot infection. There were highly significant differences 
between citranges, Yanco hybrid 3694 being more resistant and Yanco hybrid 
3700 more susceptible to collar rot than the other citranges (S.H. mean 
(nt=10)=-+0-027 on 52 degrees of freedom). The reactions of Scarlet and 
Cleopatra mandarins did not differ significantly (overall mean (n!—14)—1-66 
with S.E. +0-122 on 12 degrees of freedom). 


(3) Reaction of P. trifoliata Selections and Trifoliate ‘‘ Orange” Hybrids to P. 
citrophthora: Root Inoculation of P. trifoliata 


Seed was obtained from trees at the Horticultural Research Station, 
Somersby, where a collection of 148 clones of P. trifoliata collected in N.S.W. and 
two introduced from U.S.A. are maintained. 

Experiment 1: Ten seedlings 10 cm. tall of each of 48 clones of P. trifoliata 
and one clone of rough lemon were planted in troughs 3 ft. x2ft.x6in. 
containing steam-air treated (140° F./30 min.) soil. The soil had been inoculated 
with a mixed inoculum of five-week-old corn-meal sand cultures of 12 pathogenic 


2) 


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124 REACTION OF SEEDLINGS OF CITRUS TO PHYTOPHTHORA CITROPHTHORA 


isolates of P. citrophthora. Two months after planting, the plants were washed 
free of soil and the root damage assessed. 


Despite the severity of the test, P. trifoliata seedlings showed little root 
injury. No infection of mature tissue occurred, but root tips were often infected 
or rotted away. Of the rough lemon seedlings, nine died within one month of 
transplanting into the infected soil and the one remaining plant was surviving 
with one lateral root, the taproot, having been rotted. 


Experiment 2: One clone of rough lemon and 129 clones of P. trifoliata were 
planted in one gallon containers of steam-air treated soil (140° F./30 min.), 10 
seeds per container. Dormant seedlings were inoculated when 7 in. tall by 
adding to each tin 1 gm. of mixed corn-meal sand inoculum comprising 10 
isolates of P. citrophthora which had grown for one month. 

All P. trifoliata seedlings were highly resistant to P. citrophthora root rot. 
The rough lemon seedlings developed severe root rot. 


TABLE 4 
Reaction of Citrus spp. and Related Genera to Stem Inoculation with Phytophthora citrophthora 


Length of 
Lesion 
in Cm. 
(Mean of 10) 


Common Name Botanical Name 


Yanco hybrid 3694* Poncirus trifoliata x Citrus sinensis. F, 0-05 
generation 

Box orange 54—93-9 Severinia buaifolia (Poir.) Tenore 0-05 
Trifoliate orange 111 Poncirus trifoliata (L.) Raf. 0-08 
Australian wild lime Microcitrus australis (Blanco) Swing. 0-09 
Troyer citrange P. trifoliata xC. sinensis 0-1 

Box orange 54-91-11 Severimia buxifolia (Poir.) Tenore 0-11 
Yanco hybrid 3698 P. trifoliataxC. sinensis. F, generation 0-14 
Yanco hybrid 3693 BS 35 55 55 a fe 0-18 
Yanco hybrid 3697 AB bs a a Bs) . 0:18 
Trifoliate orange 113 Poncirus trifoliata (L.) Raf. 0-2 

Yanco hybrid 3700 P. trifoliataxC. sinensis. EF, generation 0-42 
Scarlet mandarin C. reticulata Blanco Ih > S7/ 
St. Michael sweet orange C. sinensis (L.) Osbeck 1-59 
Cleopatra mandarin (N.S.W.) C. lycopersicaeformis Hort. ex Tan. 1-74 
Rangpur lime 3234 O. limonia Osbeck 1-82 
Cummins sweet orange C. sinensis (L.) Osbeck 1-93 
Ruby blood sweet orange 3699 oo Ks Bs ae 1:96 
Viticultural nursery sweet orange ae 3 zi Be 1-99 
Maltese blood sweet orange 3206 A is No ae 2-07 
Moorland sweet orange ae 45 bsg ee 2-08 
Ruby blood sweet orange 3696 “e A a ‘ Ho 1 

Ruby blood sweet orange 3695 ss Bs ae i 2-2 

Harrison sweet orange a2 BS i aS 2-54 
Rough lemon C. jgambhirt Lush. 2-68 
Smooth Seville 3151 C. aurantium L.? 2-75 


* Refers to the accession number of the tree from which the seeds were obtained at the 
Horticultural Research Stations at Narara and Somersby. 


Experiment 3: Seeds of Troyer citrange, rough lemon and 13 clones of P. trifoliata 
were planted in rows (20 seeds per row) in troughs 3ft.x2ft.x6in. deep 
containing steam-air treated (140° F./30 min.) soil. When the seedlings were 
4in. high corn-meal sand inoculum of 12 isolates of P. citrophthora grown for 
one month was incorporated into the soil. After five months plants were washed 
free of adhering soil and the root damage was assessed. 


Only eight of 40 rough lemon seedlings survived. The remainder had 
callused lesions in the collar region or on the taproot. In the majority of plants 


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BROADBENT ET AL. 125 


at least 50°% of the root system had been destroyed and plants were surviving 
on two or three lateral roots as the taproot had been rotted. Root regeneration 
was a major factor in the survival of these plants. The trifoliate clones and 
Troyer citrange plants were all highly resistant to P. citrophthora. Some root 
tips had been attacked, but the disease had not spread to mature tissue. 


Stem and Root Inoculations of P. trifoliata Hybrids 

One hundred and nineteen F1 hybrids of Smooth Seville P. trifoliata, 130 
F1 hybrids of Ellendale mandarin x P. trifoliata and 10 rough lemon plants of 
nursery row size were subjected to root and stem inoculation as described in 
Sections (1) and (2). 

Stem inoculations did not produce collar rot lesions in hybrid plants, although 
the rough lemon seedlings developed collar rot. Two hybrids of Smooth 
Seville x P. trifoliata were highly susceptible to root rot, while the remainder 
were moderately resistant. Kllendale mandarin hybrids varied in their reaction. 
In most plants only the tips of feeder roots were rotted away, but in seven plants 
the percentage of roots rotted varied from 10 to 40, compared with rough lemon, 
where 50-70°% of roots were rotted. 


DISCUSSION 


The resistance of Poncirus trifoliata and the citranges to root and collar 
inoculation with P. citrophthora confirmed field and nursery observations. All 
clones of P. trifoliata tested, one of which was a tetraploid, were extremely 
resistant to Phytophthora root rot. This was in contrast to the variation found 
by Klotz et al. (1958a) in percentage decay of fibrous roots of trifoliate orange 
selections caused by P. citrophthora and by P. nicotianae var. parasitica. Carpenter 
and Furr (1962) also found the percentage survival of trifoliate orange selections 
to be highly variable following inoculation with P. nicotianae var. parasitica and 
postulated that the reaction of P. trifoliata and its hybrids to P. nicotianae var. 
parasitica may be related to the degree of dormancy. No evidence to support this 
hypothesis was obtained with P. citrophthora in trials in New South Wales. 
Furr, Carpenter and Hewitt (1963) considered that differences in susceptibility 
of trifoliate oranges may be due to differences between young and old root 
systems, as well as to differences in pathogenicity of the species and isolates of 
Phytophthora. This explanation cannot be applied to the behaviour of P. 
citrophthora in New South Wales, where it has been found that the immature 
tissue in the region of elongation of trifoliate orange roots is rotted but mature 
cells are not attacked (Broadbent, 1969). 


It is possible that the Australian clones of P. trifoliata have been derived 
from a small number of original introductions, which might explain the narrow 
range of variation. 


The New South Wales selection of rough lemon, Volkamer lemon and the 
South African selection of rough lemon showed similar susceptibilities to root 
and collar infections by P. citrophthora. These stocks are vigorous and adaptable, 
and suitable for lemons, but lack the resistance to root rot necessary for citrus 
replant situations. The Terrigal selections of rough lemon were slightly less 
susceptible to Phytophthora root rot than the commonly grown rough lemon 
selection, and further search within the species for a more resistant clone may be 
desirable. 


The tangelos and mandarins showed intermediate resistance to P. citrophthora 
and were more resistant than most of the sweet orange selections tested. The 
Ellendale mandarin, an Australian variety which is postulated to be a natural 
tangor (Swingle and Reece, 1967), is unsatisfactory as a stock as all its seedlings 
are gametic. 


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126 REACTION OF SEEDLINGS OF CITRUS TO PHYTOPHTHORA CITROPHTHORA 


Cleopatra mandarin is in limited use as a rootstock in Australia and overseas, 
and for mandarins has an advantage over rough lemon stocks. The rating of 
Emperor mandarin as slightly more resistant than rough lemon confirms field 
observations that seedling trees of this variety persist better in root rot situations. 

The resistance to root rot caused by P. citrophthora of four selections of 
Smooth Sevilles is in agreement with field observations and with the results 
obtained by Grimm and Garnsey (1969) using P. nicotianae var. parasitica. 
Under certain conditions Smooth Seville is highly susceptible to collar rot caused 
by P. citrophthora. The taxonomic status of the variety here called Smooth 
Seville is in doubt. It is not typical sour orange as described (Hodgson, 1967). 
A relatively high percentage of bifoliate and gametic seedlings is produced. 
Differences in susceptibility of unifoliate and bifoliate seedlings have been noted 
by Grimm and Garnsey (1969). Smooth Seville performs reasonably as a stock 
for lemons, but is unsatisfactory for oranges as this stock-scion combination 
declines due to tristeza virus (Stubbs, 1963). 

A high degree of susceptibility was shown by most sweet oranges. Sweet 
orange is commonly used as a rootstock on the neutral to alkaline, free draining 
sandy loam soils of the inland districts of New South Wales. Ruby blood 
selection 3118 and Parramatta orange selection 3443 were somewhat more 
resistant to Phytophthora root rot than most of the other sweet orange selections. 
It may be significant that the parent tree of the Parramatta strain of sweet 
orange was a 100-year-old seedling tree from Northmead (New South Wales). 
Tt was in this area, once a prosperous citrus growing district, that P. citrophthora 
caused serious damage in 1860 (Fraser, 1949). 

The reaction to Phytophthora root and collar rots of the range of species 
and varieties tested has provided a measure of the relative resistance of seedlings 
of many varieties of Citrus spp. and related genera. This can be used as a basis 
for further testing in the field as rootstocks and also provide information on 
resistance for future breeding purposes. 

The genetic constitution of the citrus host rootstock and the suitability of 
the environment are of prime importance in determining the course of 
development of Phytophthora root or collar rots in the field. In genera such 
as Poncirus, Microcitrus and Severinia, resistance is conferred by some factor 
of genetic constitution. Where resistance is not complete, the degree of damage 
caused can be influenced by age, chemical composition, succulence and vigour 
of the infected tissue (Carpenter and Furr, 1962). 

Under field conditions, environmental factors can influence both the course 
of the disease and the host. Soil characteristics, soil temperature, soil oxygen 
and moisture levels influence the development of citrus root rot (Klotz et al., 
1965; Stolzy et al., 1965a). A low supply of oxygen in soils prevents root 
erowth and regeneration, creating an unfavourable soil condition for infected 
plants to overcome the adverse effects of root decay (Stolzy et al., 1965b). Soil 
fertility, with its effect on tree vigour and root development (Fraser, 1949), 
and the effect of the scion on rootstock (Klotz et al., 1967) are also important. 


ACKNOWLEDGEMENTS 


The authors are indebted to Mr. A. KE. Roberts, Chief Biometrician, New 
South Wales Department of Agriculture, for statistical analysis of the data. 


References 
BrRoapDBENT, P., 1969.—Observations on the mode of infection of Phytopthora citrophthora in 
resistant and susceptible citrus roots. Proc. First International Citrus Symposium, 3: 
1207-1210. 
CARPENTER, J. B., and Furr, J. R., 1962.—Evaluation of tolerance to root rot caused by 
Phytophthora parasitica in seedlings of Citrus and related genera. Phytopathology, 52: 
1277-1285. 


PROCEEDINGS OF THE LINNEAN SoctmeTy or NEw SoutH Watzs, Vou. 96, Part 3 


BROADBENT ET AL. iT 


Fawcett, H. §8., 1913.—Two fungi as causal agents In gummosis of lemon trees in California. 
Phytopathology, 3: 194-195. 

————, 1923.—Gummosis of citrus. Part I. Gummosis due to Pythiacystis citrophthora. 
J. Agric. Res., 24: 191-213. 

Frasmur, L., 1942.—Phytophthora root rot of citrus. J. Aust. Inst. Ag. Sci., 8: 101-105. 

, 1949.—A gummosis disease of citrus in relation to its environment. Proc. Linn. 
Soc. N.S.W., 74: 5-14. 

Furr, J. R., CARPENTER, J. B., and Hewitt, A. A., 1963.—Breeding new varieties of citrus 
fruits and rootstocks for the South-west. J. Rio Grande Hort. Soc., 17: 90-107. 

Grimm, G. R., and GARNsEY, 8. M., 1969.—Root rot and tristeza tolerance of smooth seville orange 
from two sources. Cvtrus Industry, 50: 12. 

Grimm, G. R., and WHIDDEN, R., 1962-3.—Range of pathogenicity of Florida cultures of the 
foot rot fungus. Proc. Fla. Sta. Hort. Soc., 75: 73-74. 

Hoacnanpb, D. R., 1919.—Relation of concentration and reaction of the nutrient medium to the 
growth and absorption of the plant. J. Agric. Res., 18: 73. 

Hopeson, R. W., 1967.—Horticultural varieties of citrus. In Reuther, W., Webber, H. J., and 
Batchelor, L. D. (eds.), The Citrus Industry, 2nd ed., Vol. 1, Ch. 4, pp. 514, 551. Univ. 
Calif. Div. Agr. Sei., Berkeley. 

Krorz, L. J., Brrrers, W. P., and DE Wotrs, T. A., 1965.—Citrus rootstocks resistant to 
Phytophthora root rot. California Agriculture, 19: 10. 

, 1966.—Resistance to Phytophthora root rot. Calif. Citrograph., 51: 257-258. 

Kuotz, L. J., and Dz= Wotrs, T. A., 1965.—Tetrazolium, an indicator of extent of infection in 
Phytophthora root rot of citrus. Pl. Dis. Reptr., 49: 423. 

Kiorz, L. J.. De Wore, T. A., and Batnszs, R. C., 1969.—Resistance of trifoliate orange stocks 
to gummosis. Calif. Citrograph., 54: 259-260. 

Kiorz, L. J., DE Wore, T. A., and Po-Pine Wone, 1958a.—Influence of two varieties of citrus 
scions on the pathogenicity of three isolates of Phytophthora parasitica to sweet orange 
rootstocks. Phytopathology, 48: 520-521. 

, 19586.—Decay of fibrous roots of citrus. Phytopathology, 48: 616-622. 

Kiorz, L. J.. D—E Wotre, T. A., Moorz, P. W., and Newcoms, D. A., 1967.—Testing sweet 
orange rootstocks. Calif. Citrograph., 52: 387-388. 

Knorz, L. J., and Fawcrrt, H. §., 1930.—The relative resistance of varieties and species of 
Citrus to Pythiacystis gummosis and other bark diseases. J. Agric. Res., 41: 415-425. 
Kuorz, L. J., and Soxotorr, V. P., 1943.—The possible relation of injury and death of small 

roots to decline and collapse of citrus and avocado. Calif. Citrograph., 28: 86-87. 

Krorz, L. J., Stonzy, L. H., De Wotrs, T. A., and Szuszxrewics, T. E., 1965.—Rate of oxygen 
supply and distribution of root-rottimg fungi in soils. Sow Science, 99: 200-204. 

Navurivau, L. J.. Saannon, L. M., and Froticn, EH. F., 1958—Eureka lemon trifoliate orange 
incompatibility. Am. Soc. Hort. Sci., 72: 273. 

Rossetti, V., 1947.—Susceptibility of different species of Cztrus to some species of Phytophthora. 
Arq. Inst. Biol., 18: 97-124. 

Stonzy, L. H., Lerey, J., Knorz, L. J., and LABANAUSKAS, C. K., 1965a.—Water and aeration 
as factors in root decay of Crtrus sinensis. Phytopathology, 55: 270-275. 

Stontzy, L. H., Lerry, J., Knorz, L. J., and Dr Wotre, T. A., 1965b.—Soil aeration and 
root rotting fungi as factors in decay of citrus feeder roots. Soil Science, 99: 403-406. 

Stupss, L. L., 1963.—Tristeza-tolerant strains of sour orange. #.A.O. Plant Protection Buil.. 
1: 8-10. 

Swine se, W. T., and Re«Ec#, P. C., 1967.—The botany of Citrus and its wild relatives of the Orange 
subfamily. In Reuther, W., Webber,H..J., and Batchelor, L. D. (eds.), The Citrus Industry, 
2nd ed., Vol. 1, Ch. 3, p. 514. Univ. Calif. Div. Agr. Sci., Berkeley. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES. VoL. 96, Part 3 


AUSTRALASIAN CERATOPOGONIDAE (DIPTERA, NEMATOCERA) 


Part XV: THE GENUS ALLUAUDOMYIA KIEFFER IN AUSTRALIA AND 
NEW GUINEA 


MARGARET L. DEBENHAM 
School of Public Health and Tropical Medicine, University of Sydney 


[Accepted for publication 21st April 1971] 


Synopsis 
The Australian and New Guinea species of the genus Alluaudomyia Kieffer are revised. 
Ten new species and one new subspecies are described, Skuse’s Ceratopogon latupennis is transferred 
to this genus, and Tokunaga’s Alluaudomyia novaguineae is placed in synonymy with Alluaudomyia 
spinosipes Tokunaga. New data are recorded for some previously described species, and a key 
is provided for identification of the 25 known species found in the area. 


This study was prompted by the publication of Wirth and Delfinado’s 
comprehensive revision of the Oriental species of Alluaudomyia (1964). In 
this paper the authors state that ‘‘ only the Neotropical and Australian species 
are now poorly known, but undescribed material indicates that the genus is 
not well represented in the extreme south. Only 3 or 4 species collected in 
Australia... are known to us.’’ The present study raises the number of species 
recorded from the Australian mainland to 14, but of these four are New Guinea 
or Oriental species which have penetrated to the far north of Australia and a 
further three have thus far only been recorded from the north of the continent, 
although they are not yet known from New Guinea. Of the remaining seven 
species, only two, latipennis and alpina, appear to have a purely southern 
distribution. 


The first species of the genus to be recorded from the Australia-New Guinea 
area was described by Skuse in 1889 under the name Ceratopogon latipennis, 
but the correct generic status of this species has until now been unrecognized, 
and consequently it was omitted from Wirth and Deifinado’s list of world species. 
No further record of the genus in this area appeared until 1955, when Lee and 
Reye reported its occurrence in Queensland and New South Wales. In 1959 
a Single species was described from New Guinea by Tokunaga, and a further 
11 species were named from the same area in 1963, while one species previously 
described from Micronesia was also recorded. The present paper increases to 
25 the number of species known from the Australia-New Guinea area. Of these 
10 are new and one is a new subspecies of an Oriental species. 


Wirth and Delfinado placed the Oriental species in five groups based on 
characters of wing pattern, spermathecae, hypopygium and leg banding. These 
groups have not been followed in the present paper as some of the Australian 
species combine the characteristics of more than one group, resulting in a partial 
breakdown of the classification. At present it appears that a group system 
based on the number and form of the spermathecae and certain male genitalic 
characters may be possible, but the resulting groups would be too large and 
contain too many widely differing forms to be of any practical value. 


METHOD OF DESCRIPTION 
All new descriptions are based on slide specimens cleared in a mixture of 
one part absolute alcohol to one part creosote and mounted in balsam on 
microscope slides, except for the few cases where alcoholic or pinned specimens 
have been available for detailed description of the thoracic pattern. 


PROCEEDINGS OF THE LINNEAN SoctetTy or New SoutH WaAteEs, Vou. 96, Part 3 


MARGARET L. DEBENHAM 129 


Table of Measurements 


Measurements of newly described species are based on the holotype and, 
if available, the allotype, unless otherwise stated. Measurements of previously 
described species are taken directly from the original descriptions. Wing length 
is measured from the basal arculus to the wing tip, which gives a value approxi- 
mately one-tenth less than the measurement employed by Tokunaga, and 
allowance should be made for this. 


Morphological Terms 


See Wirth (1952) and, for modifications, my previous paper in this series 
(Part XIII, Proc. Linn. Soc. N.S.W., 94 (2): 145, 1970). For pupal characters, 
see Jones (1961). Length is measured from the vertex of the head to the tip of 
the abdomen. 


Illustrations 


Except for drawings of thoracic pattern, which are freehand, these were 
done with the aid of a graticule and squared paper. All are based on type 
specimens unless otherwise stated. In figures of femora and tibiae, the fore 
leg is uppermost. 


Location of Types 


Types of newly described species are lodged in the collection of the School 
of Public Health and Tropical Medicine, Sydney, unless otherwise stated. 
Paratypes, where available, are in the School of Public Health; Australian 
National Insect Collection, Canberra, A.C.T.; United States National Museum, 
Washington; British Museum (Natural History); B. P. Bishop Museum, 
Honolulu. 


Abbreviations 

S.P.H. & T.M... School of Public Health and Tropical Medicine, Sydney. 
A.N.I.C. .. Australian National Insect Collection, Canberra, A.C.T. 
ely ee: .. anteromarginal (tubercle). 

d.a.s.m. .. dorsal anterosubmarginal (tubercle). 

l.a.s.m. .. lateral anterosubmarginal (tubercle). 

d.p.m. .. dorsal posteromarginal (tubercle). 

open .. lateral posteromarginal (tubercle). 

V.p.m. .. ventral posteromarginal (tubercle) 


Genus ALLUAUDOMYIA Kieffer 

Alluaudomyia Kieffer, 1913, Voyage Ch. Alluaud et R. Jeannel en Afrique 
Orientale, Dipt., 7: 12; de Meillon, 1939, J. ent. Soc. S. Africa, 2: 7; 
Okada, 1942, Trans. Nat. Hist. Soc. Formosa, Jee olds Wart l 952. ‘Ann. 
ent. Soc. Amer., 45: 423; Tokunaga and Murachi, 1959, Ins. Micronesia, 
12: 352; Wirth and Delfinado, 1964, Paci. Insects, On: 599. Type species, 
by monotypy, Alluaudomyia imparunguis Kieffer. 

Neoceratopogon Malloch, 1915, Bull. Ill. St. Lab. Nat. Hist., 11: 310. Type 
Species, by original designation, Ceratopogon bellus Coquillett. 

Prionognathus Carter, Ingram and Macfie, 1921, Ann. Trop. Med. Parasit., 14: 
309. Type species, by original designation, Prionognathus marmoratus 
Carter, Ingram and Macfie. 

Thysanognathus Ingram and Macfie, 1922, Ann. Trop. Med. Parasit., 16: 244 
(nom. nov. for Prionognathus Carter, Ingram and Macfie, nec La Ferteé- 
Sénectere). 

Isoecacta Garrett, 1925, Seventy New Diptera, p.9. Type species, by monotypy, 
Isoecacta poeyi Garrett=bella (Coquillett). 


PROCEEDINGS OF THE LINNEAN SoctetTy or NEw SourH WatsEs, Vou. 96, Part 3 


130 THE GENUS ALLUAUDOMYIA KIEFFER 


Generic Diagnosis (from Wirth and Delfinado, 1964) 

Body moderately slender, not strongly hairy, usually small to medium, but 
occasionally larger, up to 2-7mm. LHyes bare or pubescent, contiguous to 
narrowly separated above. Palp 5-segmented, segment III usually with a small 
round pit on apical half bearing sensillae. Antenna 15-segmented, 9 with distal 
5 segments slightly elongated, 3 with distal 3 segments elongated, plume present. 
Legs slender, without strong spines, occasionally with spine-like hairs; hind 
first tarsal segment bearded ventrally, fourth tarsal segment of all legs cordate, 
fifth slender, unarmed. Empodium vestigial; 92 claws long and slender, simple, 
usually very unequal but sometimes subequal on one or more pairs of legs, ¢ claws 
all small and equal. Wing without microtrichia, macrotrichia usually present 
towards wing apex, sometimes extending over most of the membrane; first 
radial cell absent, R,,; thickened just before its junction with costa, costa 
ending before to beyond middle of wing, median fork petiolate. Abdomen 
moderately stout; 1 or 2 sclerotized spermathecae present, sometimes an 
aberrant third one also present; 4 genitalia with very diverse specific modi- 
fications, but ninth tergite usually elongate, ninth sternite usually short with 
caudomedian excavation, aedeagus usually with basal arch and distinct caudo- 
median stem, parameres usually separate, with a lateral, sometimes detached, 
basal arm, elongate stem, and distal armature of one or more spines, lobes or 
filaments. 


Key to Australian and New Guinea Species of ALLUAUDOMYIA 


1. Wing with more than 8 conspicuous dark spots ....................-2205-- sees eeees 2 
Wing with 3 or fewer conspicuous dark spots, these situated at end of costa, just before 
r-In Cross- vem, and On Mis sre scvals bis temas Mande tones an nae nbn s eels toleas Un ee ngeinenay Sue STS Ate 7 

2. Wing with 7 spots arranged on M,, M,, M,,,, Cu, and anal vein, as well as mottling on 
MAC ASUS) 5 ie su sijocs Ae aos Sor ae Buwhitoke Pegeleil tael etapa ragays ecto Rega oe Perce cyanea coflaty aeayie A ae ea 3 
\AVaiaee Trine, SPOOUS: SHURE! tin Cells Biss yrEll GS OM WAS oceccocdsccovoadcavgccvoo0adsb see 5 

3. g wing with 3 dark punctures on Rs before r-m cross-vein; ¢ and 9 wing without dark 
PUNCHUTESMONMR ieee Wee. Lees) A sitad. Ab ae AEE eae ete ate man: EPaasey ee fel: 1. personata n. sp. 


ey wie without dark punctures on Rs before r-m cross-vein ; 2 wing with 2 dark punctures 
on R 


4. Apdedeie with a very elongate caudal stem; spermatheca large, elongate oval, approxi- 
mately 0:12x0:08mm.; some macrotrichia of 9 wing with pale grey dots at bases ... 
B eC rausncre ho eeis PERE ee Bin erie Neen Ken Bini Abbie td bo Bln ah KD bo 48 2. annulipes australiensis n. subsp. 
Aedeagus with a short caudal stem ; spermatheca smaller, more rounded, 0-09 «0-06 mm. ; 
some macrotrichia of 2 wing with conspicuous black dots at bases .................... 
Gite ciel Teka weit de sive ne: SUV auch helt pale errenapeuaye pune eR Wc aegalctet tone en iar 3. tokunagat Wirth and Delfinado 


5. Wing with spots in cells very close to wing margin ; spot at end of costa scarcely extending 
on to membrane ; mid and hind femora with basal half to two-thirds dark ;. spermathecae 


without) diverticula): . cle Anne serait one nhc Ree ate mor ee a ee eae one 14. varia n. sp. (9) 
Wing spots in cells not very close to wing margin; spot at end of costa extending consider- 
ably on to membrane ; all femora broadly pale basally ; spermathecae with diverticula .. 6 
6. 9 with wing margin broadly infuseated, spot in cell M, elongate rectangular ; spot on vein R, 
absentiin= both. sexess 7. ait.t oe iksras aaa gentile ete tute een TOR RT Seen: 4. fumosipennis n. sp. 
© without conspicuous infuscation on wing margin, spot in cell M, small and round; both 
REDNESS Wailea, CHAE (SyoKoKy) Oral HEMI TB ooo saad ccccasscuneogdoccancucceed 5. unguistyla n. sp. 
7. Wing with a single, usually pale, spot at end of costa, without definite spot on membrane 
just before r-m cross-vein, although Rs and M may be slightly infuscated ............ 8 
Wing with a distinct spot before r-m cross-vein, sometimes also with a spot on R, .... 12 
Ber TV OSs DALE: Bie ee cue he oc. auc geisie Cesena ieee CoB siete wae ego an cn twen enema een cee ARC teT ch nets eer ENE 9 
Hiyesmpulbescomt ih. casei kos oki cuadae wut otis a ARES Ud eats EU REES Bier bor ona daeae Se abck aaa Cotes 11 
Owwessrentinelysyellowil ees Ws es, lets Ie Les dee eee mice cies 6. tmmaculata Tokunaga 
ILE as yma, joBks) Javony Chistian loon IeIACKS . osc anevncconp eos aencaobsonoaduooueronD bE 10 
10. Abdomen largely yellow; aedeagus of male plate-like, deeply bifid apically, somewhat 
inssCreala hinge fy iA, soos keaoe redo do nwiedooncooue€ 7. insulana Tokunaga and Murachi 
Abdomen dark brown dorsally, paler ventrally ; aedeagus of male arched, with a single, 
shortrapical spon’, ics:. seieitetne = eis Ueigkhs oe CeO eke eae 8. verecunda n. sp. 


1 Tn this part of the key it is assumed that the unknown 9 of personata resembles the female 
of the closely related bifurcata Wirth and Delfinado ; this assumption may prove incorrect. 


PROCEEDINGS OF THE LINNEAN Society or New Sours WaAtEs, Vou. 96, Part 3 


ne 


14. 


15. 


16. 


We 


18. 


11). 


20. 


21. 


25. 


26. 


MARGARET L. DEBENHAM Sil 


Wing of 2 without macrotrichia, but with a row of spinule-like setae on anterior edge ; 
§ aedeagus with a V-shaped apical notch, parameres hooked apically and with a long, slender 
RONCHI! PDROGETS . oS obs baccosedocoguasboodsmcogeseanugaouUDROboOe 9. papuae Tokunaga 

Mid femur with a narrow pale base, rest of basal half dark .......................... 23 
Wing of 2 with a moderate number of macrotrichia, without spmule-like setae; g aedeagus 
with a W-shaped apical notch, parameres rounded apically, apical process long, flattened 


Bin! HEROIC oo5,.5 0 Pala OOOO ow. stole vo 0.00 6 bade aot SOE mENOne, otc. 10. bifasciata Tokunaga 
. Spermatheeae with diverticula; aedeagus bell-shaped, its apex produced into a short, 
verily login joroesss (anes, UA, PAS AIO, BO) coscopoucseasugepedcogcdedesaccnec 20 
Spermathecae without diverticula ; aedeagus various, but never as above ............ 13 
. Very dark species, the legs dark brown with narrow paler bands preapically on all femora 
andesub-basallysandspreapically, ony alletibiaemey sees. esas ee 11. alpina n. sp. 
mecannorerexbemsive live pale wera. c-lacppscbsteed ae nate Ghobs ols ls sae? CYR Meg ocd matty iters bs ke spay ote & 14 


Spot at tip of R,,, extending well on to wing membrane, at least in 9 ; spermathecae elongate 
oval, the long axis parallel to the neck; ¢g with coxites long and narrow, parameres with 
depachedlaasalWemrmn Si panacea cl ene ee ee waeyee EA cha SeaLetohate srs nites tate Wl aisianss tksier ele woler de tts 15 
Spot at tip of R,,; scarcely, if at all, extending on to wing membrane ; spermathecae round 
or, if oval, long axis perpendicular to the neck; g with coxites rather short and stout, 
(ORTNANEIRSS Wwantay losisell Grams GibeKelNeCl bo op k boos Soouadocbe oon eodur opusebeundEoebenaes 17 
Mid tibia’ pale except for dark base and apex.................. 12. latupennis (Skuse) 
Mid tibia largely dark or at least with a distinct dark band on apical third ............ 16 
Humeral areas of scutum entirely pale; g parameres with a long, broad, tapering preapical 
PLOcEssearisinaawellidowvbeysteniy ry.) ). pera eke a et 13. appendiculata n. sp. 
Humeral areas pale, but with distinct dark punctations at bases of setae; gj parameres with 
a pale, relatively short setose preapical process arising within the apical loop .......... 

I ets ccc sip MER acs fo estat fo Mrdee ses a) susp awa aAe a cv Ata) Seeeris «Boao oes eT ea 14. varia n. sp. (3) 
Mid femur almost entirely brown, only narrowly pale basally and preapically ........ 

FIN oy hai Eee Pe SR cree 5 IARC Ona LER oka ahre chet ailena tet) cenuace: 15. platipyga Tokunaga 
Mid femur with a broad brown central band, a narrow brown preapical band and a brown 
CIOERSY THES ATIAIS) ANCE Pear, Cotte & eeeRRI COE Sina G co tea cinta MIAO OAD CeCe Oto cin OM A Cee ieee ois eet 18 
Aedeagus of g a simple arch with a single hook-like apical process arising from its dorsal 
surface ; parameres with stems slender, two long apical processes; 2 genital sclerotization 


simnjolle. Weslngyosel, macemeononoms (Iii, Gl) sscssescdnooscouasdccscuasue 16. reyei n. sp. 
Aedeagus with paired apical processes ; parameres with stems broad basally, a single apical 
process; 2 genital sclerotization larger, more conspicuous, U-shaped ................ 19 


Apical processes of aedeagus horn-like, not bent, broad basally and tapering to a rounded 
point ; parameres with a long, tapering apical process; 2 genital sclerotization distinctly 
U-shaped, without a heavily sclerotized process inside the loop of the “U” (Fig. 157) .. 
PTR det ee ee oc lets ioe lot tabs WA ethtuchemne hee, ee eLeAnls Se Be, 17. bicornis n. sp. 
Apical processes of aedeagus laterally bent before tip ; parameres with a very short, parallel- 
sided process; 2 genital sclerotization very conspicuous, U-shaped but very broad and 
rounded, a triangular, heavily sclerotized process within the loop of the ““U”’ (Fig. 162) 
«6:0 0-09.07 ALG OEAIO Bi © EUCRORE Ren cigeh RCS ROR a EE en gon EERE ch SER epee Mees 9, On Mira tee 18. fragmentum n. sp. 
Mid femur with a distinct dark brown preapical band ..............--.---+------::- 21 
Mid femur without a distinct dark preapical band ; if band present it is very pale ...... 24 
Sub-basal dark band of mid tibia confluent with dark base forming a broad basal band 

2, 21. Gubi oO GNA G (A snc dBs Sac cle RENAUD PREPS ITE ares RCI Citi ENE iti? Petrone wd Nee 19. petersi Tokunaga 
Sub-basal dark band of mid tibia separated from dark base by a distinet white or yellow 
loguavel Sea A dee Some ADRS BORE co Oo AORN ere eo COMM NO eS San ab 6. areca > Cy tree tee 


basalhaliror maid) femunientimely pale. 2225.4 2-25 see 12-20. astera Nokunaga 
. Two spermathecae, one with a short, straight diverticulum, the other with a long, undulate 


diverticulum ; stems of parameres slender, apices flattened, slightly bent laterally, sub- 
tineraaullene, slaeimolhy joomaech eodocnanccousbsseoscondssssacnsoge Mik amnnepsis iol 
Both spermathecae with straight diverticula; male unknown...... 22. smeei Tokunaga 


. Claws of fore and mid legs subequal in 2; diverticula of spermathecae slightly swollen 
apically ; stems of parameres very narrow on basal half, broader on apical half, apices short, 
pomped wbentrabenisbipansles|bOeStem ere) leila 23. spinosipes Tokunaga 
Claws of fore and mid legs very unequal in 2; diverticula not swollen apically ; parameres 
GUS Ty Sm ete ARE uN Aeon PPS ay SEN has SER ERMC UOS «GROEN ARE ILLS AWE os a ec ea eee, DO 
Fmd eters fuscous sonm basal! jwo-bhirds! Pele else ae eae eee 24. NEG. Now 1 
Hind femur with base broadly pale and a dark central or post-central band .......... 26 
Diverticula of spermathecae short, subequal, about 0-012 mm. long; basal half of stem of 
parameres strongly swollen, apices not undulate, somewhat rounded .. 25. brandti Tokunaga 


Diverticula of spermathecae unequal, longer, about 0-051 and 0-042 mm. long; stems of 
parameres only slightly swollen basally, apices slender, undulate, sharply tapered .... 
Solio alo b's Cid CLS SIS Oe ib CII Sie DIO totes Sib ee BloInlS Slo otelbib wo nacre MAO das Odenton A Nol an irays¥er=) 


PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES. Vou. 96, Part 3 


132 THE GENUS ALLUAUDOMYIA KIEFFER 


~ SY SS 
Qa Se 
TT 


fi: RNS RON 
Winn mie ro : 


ie 


Figs 1-3. Alluaudomyia personata. 1, g wing, X55; 2, g hypopygium, x200; 3, d 
parameres, x 200. Figs. 4-9. Alluaudomyia annulipes australiensis. 4,2 wing, x55; 5, ¢ wing 
(Townsville paratype), «55; 6, 2 spermatheca, x 200; 7, ¢ hypopygium (Townsville paratype), 
«200; 8, $ paramere (Maprik paratype), 200; 9, g coxite and style (Maprik paratype), 
«200. Figs. 10-14. Alluaudomyia tokunagai. 10, 2 wing (Siutmeri specimen), X55; 11, ¢ 
wing (Innisfail specimen), x 55 ; 12, 2 spermatheca (Korogo specimen), x 200; 13, g hypopygium 
(Innisfail specimen), «200; 14, g parameres (Korogo specimen), x 200. 


PROCEEDINGS OF THE LINNEAN SocreTy or NEw SoutH WALES, Von. 96, Part 3 


MARGARET L. DEBENHAM Lays 


1. ALLUAUDOMYIA PERSONATA n. sp. (Figs 1-3, 15-16) 
Type: Holotype 3. 


Type Locality: Moran’s Creek, Innisfail, Queensland (5.ix.1963, H. 
Standfast). 


A small, brown and yellow mottled species, very similar to A. bifurcata 
Wirth and Delfinado. Female unknown. 


Male: Length 1-28 mm., wing 0:96 x0-37 mm. 


Head brown, vertex yellow. Eyes bare, separate. Palp brown, segment III 
with a small apical pit bearing sensillae (Fig. 15). Antennal segment II brown, 
segments ITI—XI whitish, XIJ—XIII brown, rest missing ; plume brown. 


Scutum yellow with brown mottling and punctures, scutellum yellowish 
but narrowly fuscous centrally, bearing 2 setae, postscutellum narrowly yellow 
on anterior edge, rest dark brown, pleuron yellow on upper half, brown on lower 
half. Coxae and trochanters light brown; legs whitish, all femora with basal 
half dark, femora and tibiae with many brown bands (Fig. 16), hind tibial comb 
of 5-6 spines; fore and hind first tarsal segment and basal half of mid first 
segment brown, rest of tarsi fuscous. Claws of fore and mid legs missing, of 
hind legs very small, less than one-third the length of the fifth tarsal segment, 
paired and equal, bifid at tip. 


Wing (Fig. 1) with a dark spot before the r-m cross-vein, one at junction 
of costa and R,,;, extending on to the wing membrane, and 7 posteriorly, one 
near apex of M,, one near either end of M,, one near apical end of M,,,, Cu,, 
and anal vein, and one on M,,, just beyond f{MCu,, as well as 3 small punctures 
on Rs and 1 on R,. Macrotrichia restricted to the wing margin between the 
ends of the costa and M,. MHaltere with stem white, knob fuscous but dark 
brown apically. 


Abdomen very pale brown, pleural membranes grey. Hypopygium (Figs 
2-3) brown, coxites narrow and curved, each with a strongly recurved ventral 
root, styles. whitish, slightly swollen basally ; aedeagus deeply arched, with a 
v-shaped notch apically, caudal stem with a pair of slender apicolateral processes, 
parameres each with an attached basal arm, stem slightly curved, swollen 
apically, preapical process moderately long, approximately one-third the length 
of the stem. 


Distribution: Known only from the type locality. 


This species is very close, in both coloration and form of genitalia, to 
Alluaudomyia bifurcata Wirth and Delfinado from Malaya and Thailand. 
However, it can be distinguished by the dark bases on all femora, the recurved 
ventral root of the coxites, the pointed, not rounded, sides of the apical notch 
of the aedeagus, and the much longer preapical processes of the parameres. It 
can be distinguished from other similar species known from the area by the form 
of the male genitalia and the presence, in the male, of punctations on Rs; 
assuming that the female closely resembles the female of bifurcata, it will be 
distinguishable from the females of annulipes australiensis and tokunagai by the 
absence of punctations on R,,,; and the absence of dots at the trichial bases. 


2. ALLUAUDOMYIA ANNULIPES AUSTRALIENSIS n. subsp. (Figs 4-9, 17-22) 
Types: Holotype 4, allotype 9 and 26 $¢g and 9 9° paratypes. 


Type Locality : Townsville, Queensland (holotype 0001-0300 hours, allotype 
2100-2359 hours, 18.xi.1955, light trap, Belgian Gardens, flying fox bait, A. K. 
O’ Gower). Paratypes from ‘Townsville (12 33, Ors 9 exale 1955, 1900-2100 hours, 
Belgian Gardens, A. K. O’Gower; 2 gg, 1 9, 13.xi.1955, 2100-2359 hours, 
light trap, Belgian Gardens, mangrove tree, A. K. O’Gower ; le OFF 4a asl Oboe 


PROCEEDINGS OF THE LINNEAN SocreTy or NEw SoutH Wates, Vou. 96, Part 3 


134 THE GENUS ALLUAUDOMYIA KIEFFER 


light trap, Belgian Gardens, A. K. O’Gower; 1 3, 17.xi.1955, 1920-2100 hours, 
light trap, Belgian Gardens, flying fox bait, A. K. O’Gower; 4 gg, same data 
as holotype; 2 gd, same data as holotype but 0300-0530 hours ; 1 3, 1 9, same 
data as allotype; 1 g, 20.xi1.1955, 2200-0600 hours, light trap, Belgian Gardens, 
mango tree near polluted swamp, A. K. O’Gower), from Darwin, Northern 
Territory (1 9, 27—29.x1.1957, 2400-0630 hours, light trap, Maranga, H. J. Reye ; 
1 g, 1 9, 13-14.vi.1958, light trap, Quarantine Stn., H. J. Reye; 1 3, same data 
but 16-17.vi.1958 ; 1 9, same data but 4-5.vi1.1958, N. J. light trap; 1 9, same 
data but 12—13.vii.1958 ; 1 9, same data but 26—27.v1i.1958, suction light trap), 
and from Maprik,t New Guinea (1 4, 1958). 


These specimens are very close to Alluaudomyia annulipes Wirth and 
Delfinado, described from Malaya and Thailand, but the males differ markedly 
in the length of the apical process of the parameres. In the Asian specimens 
this is very short and spine-like, but in the Australian and New Guinea specimens 
it is considerably longer, being approximately one-fourth the length of the stem. 
As this feature is consistent throughout the present series, the Australian-New 
Guinea form is here regarded as a distinct subspecies, at least until such time as 
intermediate forms may be collected. The females of annulipes australiensis 
can be distinguished from females of the nominate race only by locality of 
collection. 


Male: Length 1-59 mm., wing 1-01 x0-38 mm. 


Head brown, frons yellowish. Eyes bare, just contiguous. Palp brown, 
segment III with an apical pit bearing several sensillae (Fig. 18). Antennal 
segments Ii-III brown, IV—XI very pale, XII light brown, XIII brown on 
basal half, white on apical half, XIV brown with apical fourth white, XV brown 
(Fig. 20); plume yellowish-brown. 

Scutum yellowish with brown mottling and punctures, scutellum yellow, 
with 4 setae, postscutellum brown centrally, ochreous laterally, pleuron ochreous. 
Fore coxae ochreous, mid and hind coxae brown, trochanters brown, fore pair 
paler ; femora and tibiae yellow with brown bands and punctures as figured 
(Fig. 21), number of narrow bands variable, hind tibial comb of 5 spines; tarsi 
fuscous, fore segment I, base and apex of mid segment I, all of hind segment I 
and all fifth tarsal segments light brown. Claws of all legs small, under half the 
length of the fifth tarsal segment, paired and equal. 


Wing (Fig. 5) with a spot before r-m cross-vein, one on R, (the holotype 
also has a second small spot on R,), one at junction of R,,; and costa, and 7 
small spots posteriorly, one near apex of M,, one near either end of M,, one near 
apical end of M,,, and Cu,, one at f{MCu,, and one on anal vein. Macrotrichia 
restricted to the wing margin between the ends of the costa and M,. Haltere 
whitish, apex of knob brown. 


Abdomen light brown, pleural membranes brown. Hypopygium (Figs 7-9) 
brown, the coxites yellowish apically and on their inner faces, styles bidentate, 
apices brown; aedeagus with a shallow, rounded basal arch and an elongate, 
apically forked caudal stem, parameres with a short, attached basal arm, stem 
slightly bent, apex expanded, blunt, bearing a flattened, tapering process about 
one-fourth the length of the stem. 


Female: Length 1:67 mm., wing 1:35 x0-:53 mm. 
Generally similar to male, differing as follows : 


Approximately 17 mandibular teeth. Antennal segments IV—V white, 
VI-VIII brown centrally, base and apex broadly white, IX—X brown centrally, 
base and apex light brown, XI—XTV brown with apex white, XV brown (Fig. 19). 


1 Specimens with the data ‘“‘ Maprik, New Guinea ”’ are part of bulked light trap collections 
sent to §.P.H. and T.M. for study by Dr. W. Peters. 


PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SoutH WALES, Vou. 96, Part 3 


MARGARET L. DEBENHAM 135 


Claws of all legs unequal, ratio of length of claws to fifth tarsal segment 
24:12: 25 in fore, 23: 12: 24 in mid, 17: 7: 20 in hind (Fig. 22). 

Wing (Fig. 4) also with several small brown punctures on Rs and R,,;. 
Macrotrichia more extensive, on apical half of wing and in anal cell, with greyish 
dots at bases of many macrotrichia. 

Cerci pale fuscous. Spermatheca large, elongate oval, with hyaline punctures 
anteriorly (Fig. 6). 

Distribution: New Guinea, Northern Territory, north-eastern Queensland. 

This subspecies can be distinguished from tokunagai by the form of the 
male genitalia, the larger, more elongate oval spermatheca, and the paler, less 
conspicuous dots at the trichial bases on the female wing. 


3. ALLUAUDOMYIA TOKUNAGAI Wirth and Delfinado. (Figs 10-14, 23-27) 


Alluaudomyia splendida Tokunaga, 1963 (nec Winnertz, 1852), Pacif. Insects, 
5: 216 (Type locality: Keravat, New Britain. Allotype from Goldie 
River, nr. Port Moresby, New Guinea; paratypes from Minj and Lae, 
New Guinea.) 

Alluaudomyia tokunagat Wirth and Delfinado, 1964, Pacif. Insects, 6: 633. 
Nom. nov. for splendida Tokunaga preoce. Winnertz, 1852. 

Specimens examined: New Guinea: Korogo, Sepik River (2 gd, 1 9, 
8.i11.1964, D. H. Colless) ; Siutmeri, Sepik River (2 929, 17.iv.1964, D. H. Colless). 
Northern Territory: Darwin (1 g, 26—27.vii.1958, Quarantine Stn., suction 
light trap, E. J. Reye). Queensland: Innisfail (2 ¢g, 13.vi.1963, Eubenangee 
Swamp, H. Standfast; 1 3, 5.ix.1963, Moran’s Creek, H. Standfast). 

Characteristics: Medium-sized mottled species. Head yellowish-brown, 
vertex with dark median spot, eyes bare. Scutum yellow, mottled with brown 
dots and stripes, scutellum entirely yellow or with a narrow central fuscous band, 
4 setae in female, 2 in male, postscutellum yellow ; legs yellow with many brown 
bands, basically as figured (Fig. 26) but number of narrow bands variable, 
female claws all very unequal, ratio of length of claws to fifth tarsal segment 
Zo lao) 27 in fore) 29 212-5 228-5 im mid, 23-5: 9-5) 225 in hind (Fig. 27), 
male claws all small and equal. Female wing with dark spots on Rs, R,, R,,;, 
M,, M,, Ms,,, Cu,, and anal veins, and a spot proximal to r-m cross-vein, also 
with dark dots at some trichial bases (Fig. 10), male wing lacking spots on Rs 
and at trichial bases (Fig. 11), female with a moderate number of macrotrichia 
on apical half of wing and a few in anal cell, male with macrotrichia restricted 
to anterior edge and apex. MHaltere with all or part of knob, and sometimes 
part of stem, fuscous. Abdomen pale ochreous to pale brown. Spermatheca 
(Fig. 12) large, single, oval, with hyaline dots before base of duct. Aedeagus 
with a deep basal arch and a small, four-pointed apical lobe, parameres separate, 
apical third expanded and flattened, with a slender preapical process (Figs 13-14). 

Distribution: New Britain, New Guinea, Northern Territory, north-eastern 
Queensland. 

This species is readily distinguished from annulipes australiensis by the 
form of the male genitalia, the smaller, more rounded spermatheca and the very 
dark trichial bases on the female wing. 


4, ALLUAUDOMYIA FUMOSIPENNIS n. sp. (Figs 28-49, 46-51) 

Types: Holotype Q, allotype 3, 20 gg and 29 99 paratypes. 

Type Locality: Darwin, Northern Territory (holotype 14—15.vi.1958, 
allotype 30-31.x.1957, Quarantine Stn., N.J. light trap, E. J. Reye). Paratypes 
from Darwin (2 $3, 2 99, 27—28.vi.1956, 1 g, 22—23.xi1.1957, 1 g, 10—11.vi.1958, 
1 9, 11-12.vi.1958, 3 G9, 13-14.vi.1958, 1 9, same data as holotype, 1d, 299, 


PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SoutH WaAtzEs, Vou. 96, Part 3 


136 THE GENUS ALLUAUDOMYIA KIEFFER 


17 


Figs 15-16. Alluaudomyia personata. 15, 3 maxillary palp, segments III-V, x 350; 
16, g femora and tibiae, «90. Figs 17-22. Alluaudomyia annulipes australiensis. 17, ° 
maxillary palp, x 350; 18, g maxillary palp, x 350; 19, 2 antennal segments VITI-XV, x 350; 
20, g antennal segments XI-XV, «350; 21, g femora and tibiae, x90; 22, 2 fore (right) and 
hind (left) tarsus V and claw, «350. Figs 23-27. Alluaudomyia tokunagai. 23, 2 maxillary 
palp (Siutmeri specimen), x 350; 24, 9 antennal segments VIII—XV (Siutmeri specimen), x 350: 
25, g antennal segments XI—XV (Korogo specimen), « 350; 26, 2 femora and tibiae (Siutmeri 
specimen), x90; 27, 2 fore (r7ght—Siutmeri specimen) and hind (left—Korogo specimen) tarsus V 
and claw, x 350. 


PROCEEDINGS OF THE LINNEAN SocrETy oF NEw SoutH WatEs, Vou. 96, Part 3 


MARGARET L. DEBENHAM 137 


NOR oS re} te) os AD) 
W ° SMG 0% og a 


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Nee Noe 
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Jpeg? Qe iy 36 
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ia 
ONE NT Es Ay EN 
BONA LAU wD 


Figs 28-40. Alluaudomyia fumosipennis. 28, dorsum of thorax (Mt. Crosby specimen) ; 
29, 2 wing, x55; 30, g wing, «55; 31, 2 spermathecae, «200; 32, ¢ hypopygium, 
<x 200; 33, g parameres, x 200: 34, jg coxite and stvle. x 200; 35, g style (paratype), 
x 200; 36-40, pupa. 36, operculum, x 200; 37, dorsal tubercles of cephalothorax, x 200: 
38, tubercles of 7th abdominal segmeat, 200; 39, tubercles of 8th abdominal segment, 
x 200; 40, anal segment, «200. Figs 41-45. Alluaudomyia unguistyla. 41, 2 wing, x55; 
42, g wing (Noonameena specimen), «55; 438, 2 spermathecae, x 200; 44, g hypopygium 
(Yeerongpilly specimen), x 200; 45, g parameres (Yeerongpilly specimen), x 200. 


PROCEEDINGS OF THE LINNEAN Soctety oF New SourH WALES, Von. 96, Part 3 


138 THE GENUS ALLUAUDOMYIA KIEFFER 


18-19.vi.1958, 1 g, 22—-23.vi.1958, 3 gd, 19, 28-29.vi.1958, 1 9, 2-3.vii.1958, 
1 Go, 25 6. varll9o8) OO Se vaiel 9585 2 OOS a 2s aval Oa eee 
12 99, 26—27.vii.1958, Quarantine Stn., light trap, EH. J. Reye; 1 g, 26-27.xi.1957, 
1 3g, 1-2.vili.1958, Kast Arm Convent, light trap, E. J. Reye; 1 9, 21.iv.1958, 
1 9, 5-6.v.1958, N.J. light trap, J. Dyer; 1 g, 4—5.vii.1958, Winellie, suct. light 
trap, E. J. Reye). 

A small, mottled species, the wing with many spots on the veins and 
membrane. 

Female: “Length 1-36 mm., wing 0-82 «0-42 mm. 

Head brown, vertex and frons ochreous. Hyes bare, contiguous. Mandible 
with 11 teeth. Palp pale brown, segment III with a shallow preapical pit bearing 
several long sensillae (Fig. 46). Antennal segment IT ochreous, III-IX brown 
with bases whitish, X-—XV brown (Fig. 47). 

Seutum light brown, humeral areas yellow, the whole with a pattern of 
mid and dark brown (Fig. 28), scutellum yellow with a brown band each side of 
centre, with 4 setae, postscutellum dark brown centrally, paler laterally, pleuron 
yellowish with a brown transverse central band. Coxae and _ trochanters 
yellowish-fuscous ; femora and tibiae whitish, the fore femur with a broad 
central brown band and brown apex, mid femur with a broad brown sub-basal 
band, narrow brown preapical band and brown apex, hind femur similar to fore 
femur, fore tibia with brown base and apex and a broad brown central band, 
mid tibia with brown base and apex, a narrow brown sub-basal band and a broader 
preapical band, hind tibia similar to fore but central band narrower (Fig. 49), 
hind tibial comb of 6 spines; tarsi whitish but hind first tarsal segment pale 
brown, segments IV and V of all legs pale fuscous. Claws of all legs unequal, 
ratio of length of claws to fifth tarsal segment 16:11:19 in fore, 18:11:18 
in mid, ?: 8:16 in hind (13: 7:16 in paratype) (Fig. 50). 

Wing (Fig. 29) with extensive dark markings, wing margin from beyond 
end of costa to end of M,,, shaded grey. Macrotrichia numerous on apical half 
of wing. Haltere white with apex of knob brown. 


Abdomen pale brown, darker distally. Cerci white. Spermathecae (Fig. 31) 
two unequal, subspherical, each with a short diverticulum. 


Male: Length 1-44mm., wing 0-78 x 0-30 mm. 
Generally similar to female, differing as follows : 


Antennal segment II dark brown, flagellum light brown (Fig. 48); plume 
brown. 


Claws of all legs small, equal, simple, just over half the length of the fifth 
tarsal segment. 

Wing (Fig. 30) without dark margin, spotting less extensive. Macrotrichia 
restricted to the wing margin between the ends of the costa and M,. Haltere 
white. 


Abdominal segments brown with posterior edges broadly white, the anterior 
three segments largely white. Hypopygium (Figs 32-34) brown, coxites rather 
short, styles yellow-brown, long, strongly curved and sharply pointed ; aedeagus 
with basal arch extending to about half total height, apex bent ventrally, 
parameres short, stems swollen centrally, contiguous on apical half, apices 
eurved laterally. 


Pupa (Gilruth Plains specimen): Light yellowish-brown, slightly darker on 
dorsum of thorax. Respiratory trumpet short, about four times as long as 
broad, with about 9 pairs of spiracles on apical half (Fig. 51). Operculum 
short and broad, median tubercle not distinguishable, a.m. tubercles situated 
between lateral corners, each with a short, fine spine, remaining surface without 


PROCEEDINGS OF THE LINNEAN SoctETY or New SoutH WALES, Vou. 96, Part 3 


MARGARET L. DEBENHAM 139 


spines, covered almost entirely with small tubercles (Fig. 36). Dorsal tubercles 
of cephalothorax as figured, 1, 2 and 3 with a short spine, 4 with a seta of medium 
length, 5 with pore only (Fig. 37). Tubercles of abdominal segments 3-7 as 
figured, d.a.s.m. 2 and l.a.s.m. with a short spine, d.a.s.m. 1 with a long, fine 
spine, all five d.p.m.’s present, 2, 3 and 5 with a short spine, 1 and 4 appear to be 
without spines; 1.p.m.’s 1 and 3 large, with a short spine, 2 similar but with a 
long fine spine, v.p.m.’s 1 and 3 with a short spine, 2 with a long spine (Fig. 38). 
Abdominal segment 8 lacking d.a.s.m.’s, la.s.m. and d.p.m.’s 3-5 (7), d.p.m. 2, 
l.p.m.’s 1 and 3 and v.p.m.’s 1 and 3 with a short spine, d.p.m. 1, l.p.m. 2 and 
v.p.m. 2 with a long, fine spine (Fig. 39). Anal segment with a basal band of 
2-3 rows of small spines (Fig. 40). 

Additional specimens: Queensland: Cloncurry River (19, pinned, 1.v.1955, 
1715-1740 hours, Fort Constantine, net along damp edges of river bed, E. J. 
Reye) ; Longreach (1 9, 23.iv.1955, ‘* Leander ”’, net on pond shore, E. J. Reye) ; 
Merivale Ck.-Maranoa R. junction (1 g, 1 9, 6.xi.1952, 6-9 p.m., suction light 
trap, A. L. Dyce); Mt. Crosby (1 3, 6 29, 9’s in alcohol, 5.xii.1965, dusk-dawn, 
Lake Manchester Rd., 250 yds. from Brisbane Rd., in narrow, well-grassed, 
timbered gully, light trap, A. L. Dyce and M. D. Murray); Redbank Plains 
(1 9, 12.xii.1967, light trap, A. L. Dyce) ; Gilruth (1 9,1 9, 26.vii.1963, light trap, 
A. L. Dyce and M. D. Murray ; 3 gd, 5 99, 15.xii.1963, light trap, A. L. Dyce) ; 
Gilruth Plains (1 3, 2.11.1963, light trap, 2 99, 1.111.1963, bred, A. L. Dyce and 
M. D. Murray); Goodar Crossing, Weir R. (2 99, 28.111.1953, 1730-2130 hours, 
suct. light trap, W. E. Poole); Yelarbon (1 4, 2 99, 27.11.1953, 1800-2115 hours, 
4 OO, 27—28.111.1953, 2115-0045 hours, suct. light trap, W. E. Poole) ; Noondoo 
(7 29, 29-30.111.1952, 2145-0130 hours, bored spring, suct. light trap, W. E. 
Poole; 1 9, 16.xii.1963, light trap, A. L. Dyce); Texas Stn. (1 4, 26.11.1953, 
dusk—2130 hours, suction light trap, A. L. Dyce). New South Wales: Moree 
(1 g, 1 9, 1951, light trap, A. L. Dyce; 1 3, no data except collector, A. L. Dyce, 
but probably from Moree ; 1 9, 30.1.1952-31.1.1952, 2030—dawn, mercury vapour 
light trap, A. L. Dyce; 5 gd, 23.11.1963, to light, A. L. Dyce and M. D. Murray) ; 
Mungie Bundie, Meehi R., Moree (1 3, 16.11.1952, 2000-2130 hours, suct. light 
trap, A. lL. Dyce); Bundy Creek, via Moree (1 9, 1.xi.1951, light trap, 7 gd, 
6.xi1.1951, 1700-2000 hours, 10 34, 6.xii.1951, 1800-2100 hours, 15 gg, 5 99, 


il é gynandromorph, 7.xii.1951, 2400-0200 hours, 13 gg, 6 99, 7.xii.1951, 


0200-0500, 1 g, 7.xii.1951, 0500-0700 hours, mercury vapour light trap, A. h. 
Dyce; 1 9, pinned, v.1952, 2 99, 20.v.1952, 1600 hours, net in creek bed, 1 9, 
7.vi.1952, 1600 hours, net in creek bed, E. J. Reye); Hornsby (1 dg, 5.1.1957, 
1 g, 9.11957, light trap, D. J. Lee); Glenfield (29.111.1957, light trap, Keast). 


Distribution: Northern Territory, central and south-eastern Queensland, 
eastern New South Wales. 


This widespread species can be distinguished from the similar wngwistyla 
n. sp. by the absence of a spot on wing vein R,, the presence of two well-developed 
distal spots in cell R,;, and the elongated spot in cell M,. In addition, the grey 
wing Margin in the female is very conspicuous, the costa in the female never 
reaches the midpoint of the wing, knee joints are never pale, and the parameres 
of the male tend to be shorter and very strongly swollen centrally. 


The shape of the style in the male of this species varies, in some specimens 
being very strongly curved basally, as in the allotype, in others (as in the paratype 
figured, Fig. 35) being much more gently curved. In the type series the latter 
form predominates, but among the other specimens the former is most common. 


5. ALLUAUDOMYIA UNGUISTYLA n. sp. (Figs 41-45, 52-57) 
Types: Holotype Q, allotype ¢ and 1 2 paratype. 
Type Locality: Careel Bay, New South Wales (20.11.1958, light trap, D. J. 


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140 THE GENUS ALLUAUDOMYIA KIEFFER 


Lee). Allotype from Newport, New South Wales (12.xii.1956, light trap, W. 
Wirth), paratype from Careel Bay (10.iv.1958, light trap, D. J. Lee). 


A medium-sized, mottled species, very similar to fumosipennis n. sp. 
Female: Length 1-70 mm., wing 1-01 x0-50 mm. 


Head brown, vertex light brown, frons ochreous. Eyes bare, contiguous. 
Mandible with 11-12 teeth. Palpal segments I-II white, III-V pale brown, 
segment III with a small, shallow preapical pit bearing sensillae (Fig. 52). 
Antennal segment IT brown, segments ITI-IX light brown with bases broadly 
paler, X-—XV brown (Fig. 53). 


Scutum brown, paler laterally, humeral areas yellow with two small brown 
dots, dorsum with darker brown pattern and dark dots at setal bases, pattern 
similar to that of fuwmosipennis, scutellum pale yellow but very broadly dark 
centrally, with 4 setae, postscutellum brown, darker centrally, pleuron light 
brown to yellowish-brown with a dark central transverse band. Fore coxae and 
all trochanters yellowish-fuscous, mid and hind coxae brown ; femora and tibiae 
whitish, the fore femur with a broad brown central band and brown apex, the 
mid femur with a broad precentral brown band, a narrow preapical brown band 
and a brown apex, hind femur as in fore but central band narrower, fore tibia 
with base very narrowly yellowish, narrow sub-basal brown band, broad central 
brown band and brown apex, mid tibia with base and apex as in fore, a narrow 
brown band at basal third and one at apical third, hind tibia as in fore but 
central band narrower (Fig. 55), hind tibial comb of 7-8 spines; tarsi whitish 
except hind first tarsal segment brown, distal two segments of all legs fuscous. 
Claws of all legs unequal, ratio of length of claws to fifth tarsal segment 16 : 7:17 
in fore and mid, 13: 4:16 in hind (Fig. 57). 


Wing (Fig. 41) with extensive dark markings, pattern similar to fumosipennis, 
but a dark spot present on R,, one spot in cell KR; much reduced, spot in cell M, 
not elongated, wing margin not shaded grey. Macrotrichia numerous on apical 
half of wing. Haltere white. 


Abdomen brown. Cerci white. Spermathecae (Fig. 43) two, subequal, 
subspherical, each with a short diverticulum. 


Male: Length 1-71 mm., wing 1:10 x0-36 mm. 


Generally similar to female, differing as follows: Antennal segments 
III-XI whitish, XII pale basally, brown apically, XITI-XV brown (Fig. 54) ; 
plume brown. 


Pale bases of tibiae more distinct, femora also narrowly yellow apically, 
so knees conspicuously pale. Claws of all legs equal, simple, half the length of 
the fifth tarsal segment. 


Wing (Fig. 42) with central spot in cell R, entirely absent. Macrotrichia 
restricted to wing margin between the ends of the costa and M,. 


Abdominal tergites brown with posterior edges white, sternites whitish. 
Hypopygium (Figs 44—45) brown, very similar in appearance to that of fwmo- 
sipennis, but the parameres tend to be longer and not as conspicuously swollen 
centrally. 


Additional specimens: Queensland: Yeerongpilly (1 ¢, 1950, light trap, 
R. Riek; 1 9, 4.viii.1951, light trap, E. J. Reye); Moggill (1 9, 4—5.xi.1965, 
G. L. Cannon); Dinmore (1 9, 5.xii.1965, light trap, A. L. Dyce and M. D. 
Murray) ; Coombabah Creek, nr. Pacific Highway (1 9, 13.v.1953, 1630 hours, 
net, E. J. Reye). New South Wales: ‘‘ Noonameena’”’ Stn. via Bingara (1 9, 
24.1x.1952, swept in sunlight from above pool, A. L. Dyce; 1 4, 20-24.x.1952, 
A. Ll. Dyce); Hornsby (1 9, 24.1%.1956, 1 g)24:x.1956, 1 9, 29:x.1957, light 
trap, D. J. Lee). 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Watgss, Vow. 96, Part 3 


MARGARET L. DEBENHAM 141 


Figs 46-51. Alluaudomyia fumosipennis. 46, 2 maxillary palp, x350; 47, 2 antennal 
segments VITI-XV, x 350; 48, ¢ antennal segments XI-XV, x 350; 49, 2 femora and tibiae, 
x90; 50, 2 fore (right) and hind (left) tarsus V and claw, x 350; 51, pupal respiratory trumpet, 
x350. Figs 52-57. Alluaudomyia unguistyle. 52, 2 maxillary palp, x 350; 53, 2 antennal 
segments VIII-XV, «350; 54, g antennal segments XI-XV, x 350; 55, 9 femora and tibiae, 
x90; 56, 2 femora and tibiae (Moggill specimen), x90; 57, 2 fore (wpper) and hind (lower) 
tarsus V and claw (Moggill specimen), x 350. 


PROCEEDINGS OF THE LINNEAN SocreTy or NEw SoutTH WaALEs, VoL. 96, Part 3 


142 THE GENUS ALLUAUDOMYIA KIEFFER 


Distribution : South-eastern Queensland, eastern New South Wales. 

This species is very close to fumosipennis n. sp., but it can be differentiated 
by the presence of a dark spot on wing vein R, in both sexes, the smaller, more 
rounded spot in cell M,, and the reduction of one of the spots in cell R,, in the 
female this consisting of at most a small, irregular, linear spot, the male having 
only a very pale, minute spot, or lacking it entirely. Other characters which 
may be useful, although less reliable than the preceding, are: much less con- 
spicuous, sometimes absent, grey margin on wing than in fuwmosipennis ; knee 
joints usually conspicuously pale, but sometimes only slightly so (never pale in 
fumosipennis) ; costa in female reaching to or beyond midpoint of wing (average 
costal ratio of male also greater than in male of fumosipennis, but there is a slight 
overlap at the extremes of the ranges); parameres of male tend to be longer 
and less strongly swollen centrally. 


6. ALLUAUDOMYIA IMMACULATA Tokunaga 
Alluaudomyia immaculata Tokunaga, 1963, Pacif. Insects, 5: 219 (¢ only). 


(Type locality: Kampong Landbouw, Biak, New Guinea.) 


Characteristics : A small, pale ochreous species. Head ochreous, eyes bare. 
Scutum yellow on anterior half, with white humeral areas, ochreous on posterior 
half, scutellum yellow, with 2 setae, postscutellum ochreous; legs yellowish, 
only hind first tarsal segment slightly fuscous, claws all small and equal. Wing 
with a single pale ochreous spot at the junction of the costa and R,,;, macro- 
trichia absent. Haltere white. Abdomen very pale brown, hypopygium brown. 
Aedeagus triangular, with a very shallow basal arch, parameres separate, each 
with a detached basal arm, apex extended into a long, flattened, tapering process. 

Distribution: Known only from the type locality. 


Distinguishable from verecunda n.sp. by the pale thorax and abdomen, 
entirely yellow femora and tibiae, and form of the genitalia. In coloration it is 
closest to bifasciata Tokunaga, but it is separated by the absence of fuscous areas 
on the femora and tibiae, the bare eyes, and the form of the genitalia. 


7. ALLUAUDOMYIA INSULANA Tokunaga and Murachi 
Alluaudomyia insulana Tokunaga and Murachi, 1959, Ins. Micronesia, 12: 358. 
(Type locality: Wena I., Truk, Caroline Is. Allotype from Mt. Unibot, 
Ton I., Truk; paratypes from Mwot and Mutunlik, Kusaie, Caroline Is.) ; 
Tokunaga, 1963, Pacif. Insects, 5: 222 (New Britain). 


Characteristics : A small species. Head brown, eyes bare. Scutum brown 
with yellow and fuscous markings, scutellum yellowish, brown centrally, with 
2 setae ; legs yellowish with variable pale brown markings ; femora brown on 
basal two-thirds to three-fourths, or entirely pale brown except for yellow base, 
or with broad fuscous central bands, fore tibia pale brown on apical two-thirds 
to three-fourths, sometimes with a narrow yellow preapical band, or with a broad 
fuscous median band, mid tibia pale brown with apex dark and brownish clouds 
at basal and apical third, or pale brown with a broad basal and narrow preapical 
pale ring, or entirely yellow, hind tibia pale brown with brown cloud before 
middle and brown apex, or yellow with a broad brown central band and dark 
apex, or yellow with a small sub-basal fuscous spot and dark apex, female claws 
unequal, ratio of length of claws to fifth tarsal segment 18:17: 9, male claws 
all small and equal. Wing with dark spots at junction of costa and R,,, and 
on R,, distal parts of Rs and M before r-m cross-veins fuscous, female with a 
moderate number of macrotrichia on anterior and apical areas of wing, male 
with macrotrichia sparsely arranged along anterior margin. Haltere pale yellow 
to white. Abdomen mainly yellow, female with three small brown clouds on 
tergite III, tergites IV and V with anterior margin brown, tergite VIII brown on 


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MARGARET L. DEBENHAM 143 


anterior half, male with tergites II-V pale brown with a pair of yellowish spots. 
Spermathecae two, unequal, slightly pyriform. Aedeagus plate-like, deeply 
excavated apically giving it the appearance of a fish-tail, parameres with broad, 
expanded basal process, stems slightly tapering apically, apices bent at right 
angles, sharply pointed and with a small barb. 


Distribution: Caroline Islands, New Britain. 


The pale abdomen and distinctive genitalia identify this species. 


8. ALLUAUDOMYIA VERECUNDA n.sp. (Figs 58-60, 68-70) 
Types: Holotype g, 1 3 paratype. 


Type Locality : Maprik, New Guinea (1958). 


A small species with only a single conspicuous wing spot, but legs distinctly 
banded. 


Male: Length 1-37 mm., wing 0-71 x0-28 mm. 


Head brown. Eyes bare, just contiguous. Palp very pale brown, segment 
IIL stout, with a shallow pit just above centre bearing several sensillae (Fig. 68). 
Antennal segment II dark brown, flagellar segments lighter brown (Fig. 69) ; 
plume dark brown. 


Seutum brown, with yellowish areas anteriorly and laterally, scutellum 
yellow but brown centrally, with 2 setae, postscutellum brown, darker centrally, 
pleuron brown. Coxae brown, trochanters pale brown ; legs pale fuscous yellow, 
apices of femora and tibiae, sub-basal two-thirds of femora, apical two-thirds of 
fore tibia and broad central area of hind tibia light brown (Fig. 70), hind tibial 
comb of 4—5 spines; tarsi whitish but hind segment I brown, base of fore 
segment I and all fourth and fifth segments fuscous. Claws of all legs small, 
equal, half the length of the fifth tarsal segment. 


Wing (Fig. 58) with one light brown spot at junction of costa and R,,;, 
extending slightly on to membrane, parts of Rs, M, M, and M, streaked with 
brown. A single row of macrotrichia on the wing margin between the ends 
of the costa and M,. Haltere yellowish-white. 


Abdominal tergites dark brown, sternites paler brown, pleuron greyish. 
Hypopygium (Figs 59-60) dark brown, coxites broad basally, narrow apically, 
styles pale, short, gently curved ; aedeagus deeply arched, with a pair of preapical 
lateral folds and a short, ventrally bent caudal process, parameres separate, 
each with a short detached basal arm, base of stem expanded and curved, stem 
sinuous, apical process very long and tapered. 


Female: A female specimen from the Bariji River Valley, New Guinea 
(2,000 ft., nr. Toma, S.S.W. of Popondetta, ix.1964, R. Pullen) resembles the 
type specimens but is darker, with the brown leg bands much more conspicuous, 
and the preapical pale bands of the femora narrower. The hind claw is single 
with a short basal tooth (other claws missing), and the spermathecae lack 
diverticula. However, as this specimen is badly damaged, it seems advisable 
to await the collection of further specimens before describing the female. 


Distribution: New Guinea. 


This species is readily distinguished from immaculata Tokunaga by its much 
darker coloration, banded legs, and the form of the male genitalia. The bare 
eyes, different pattern of leg banding and form of the genitalia separate it from 
the similarly dark-coloured papwae Tokunaga, and the genitalia and dark 
abdomen differentiate it from insulana Tokunaga and Murachi. 


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144 THE GENUS ALLUAUDOMYIA KIEFFER 


Figs 58-60. Alluaudomyia verecunda. 58, wing, x90; 59, 3 hypopygium, x 350; 60, 3 
parameres, 350. Figs 61-64. Alluaudomyia papuae. 61, 2 wing (Maprik specimen), x90; 
62, 2 spermathecae (Maprik specimen), x 350; 63, 3 hypopygium (Maprik specimen), x 350 ; 
64, g parameres (Maprik specimen), x 350. Figs 65-67. Alluaudomyia bifasciata. 65, g wing 


(Musgrave R. specimen), x90; 66, 3 hypopygium (Musgrave R. specimen), x 350 ; 67, 3 
parameres (Musgrave R. specimen), x 350. 


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MARGARET L. DEBENHAM 145 


9. ALLUAUDOMYIA PAPUAE Tokunaga (Figs 61-64, 71-74) 
Alluaudomyia papuae Tokunaga, 1963, Pacif. Insects, 5: 217. (Type locality : 

Keravat, New Britain. Allotype from Maprik, New Guinea; paratype from 

Bainyik, New Guinea.) 

Specimens examined: New Guinea: Maprik (11 3g, 7 29, 1958; 1 g, 2 29, 
iv.1958; 6 93, 2 29, 6.vii.1958); Nineia, Morobe District (2 99, 10.v.1960, 
aspirated from rock ledges, Sowat R., B. McMillan). 

Characteristics: A small species with wings unmarked except for a pale 
yellowish spot at the end of the costa. Head dark brown, frons and proboscis 
paler, eyes pubescent. Thorax almost entirely dark brown but scutellum yellow 


i 
| 


f 


Figs 68-70. Alluaudomyia verecunda. 68, 3 maxillary palp (paratype), x 350; 69, g antennal 
segments XI—-XV, x 350; 70, ¢ femora and tibiae, x90. Figs 70-74. Alluaudomyia papuae. 
71, 2 maxillary palp (Maprik specimen), x 350; 72, 2 antennal segments VIIJ-XV (Maprik 
specimen), x 350; 73, 9 femora and tibiae (Maprik specimen), x 90; 74, 2 fore (72ght) and 
hind (left) tarsus V and claw (Maprik specimen), x 350. Figs 75-77. Alluaudomyia bifasciata. 
75, ¢ maxillary palp (Musgrave R. specimen), x 350; 76, g antennal segments XI-XIV (Musgrave 
R. specimen), x 350; 77, ¢ femora and tibiae (Musgrave R. specimen), x 90. 


~ 


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146 THE GENUS ALLUAUDOMYIA KIEFFER 


with a dark median spot and 2 setae; legs yellowish, with fuscous to brown 
clouds, markings more pronounced in female (Fig. 73), female claws (Fig. 74) 
unequal, ratio of length of claws to fifth tarsal segment 20-7:12-7:19-7 in 
fore, 24:15-7: 20-7 in mid, 14-7: 9-3:18-3 in hind, male claws all small and 
equal. Wings very pale, with a single very pale yellowish spot over the end of 
the costa, female without macrotrichia but with a row of spinulose setae on the 
anterior edge (Fig. 61), male with a few macrotrichia on the anterior edge. 
Haltere white. Abdominal tergites mainly dark brown, but II white, VI-VII 
fuscous, and in female posterior half of V and sometimes anterior half of VIII 
fuscous, sternites pale. Spermathecae (Fig. 62) two, brown, spherical or sub- 
spherical, subequal, sometimes also a third, vestigial, round. Aedeagus large, 
strongly sclerotized, with a V-shaped apical notch, parameres separate, expanded 
centrally, with a hook-like apex and a slender, tapering subapical process 
(Figs 63-64). 
Distribution: New Britain, New Guinea. 


This species is distinguished from the allied species bifasciata by its darker 
coloration, spinulose setae on the female wing, and form of the male genitalia. 


10. ALLUAUDOMYIA BIFASCIATA Tokunaga. (Figs 65-67, 75-77) 


Alluaudomyia bifasciata Tokunaga, 1963, Pacif. Insects, 5: 220. (Type locality : 
Keravat, New Britain. Allotype from Waris, West Irian.) 


Specimen examined: New Guinea: Musgrave R. nr. Port Moresby (1 4, 
25.11.1964, D. H. Colless). Allotype ¢ also examined. 


Characteristics: Small, pale brown to ochreous species, wing with a single 
pale spot at the end of the costa. Head pale ochreous, eyes pubescent. Thorax 
pale brown or ochreous with humeral areas yellowish, scutellum pale yellow 
with a fuscous median spot and 2 setae; legs yellow, mid femur slightly fuscous 
sub-basally and/or apically, hind femur fuscous apically (Fig. 77). Female 
claws unequal, ratio of length of claws to fifth tarsal segment 19-5: 12-5: 21-5 
in fore, 20-5: 12: 20-5 in mid, 16-5: 7-5: 20 in hind, male claws all small and 
equal. Wing (Fig. 65) with a single pale ochreous spot at the end of the costa, 
female with a moderate number of macrotrichia at the wing apex, male with 
only a few macrotrichia. Haltere white. Abdomen of female with segments 
J-II and V—VI mainly white, III-IV and VII-IX mainly fuscous, but anterior 
part of V fuscous and anterior part of VII white, abdomen of male with I ochreous 
or pale brown, II white to yellow, III-IV or V and VITI-IX brown, V or VI-VII 
pale yellow. Spermathecae two, round, equal. Aedeagus large, with a 
W-shaped apical excavation, parameres separate, each with a flattened, tapering 
apicolateral process (Figs 66-67). 


Distribution: New Britain, West Irian, New Guinea. 


The Musgrave R. specimen lacks the strong spines on the coxite which are 
present in the allotype, but is otherwise similar. The costal ratio of the allotype 
is 0-53, not 0-43 as recorded in the original description. 


This species is allied to papuae Tokunaga, but is distinguished by its paler 
coloration, absence of spinulose setae and presence of macrotrichia on the female 
wing, and form of the male genitalia. 


11. ALLUAUDOMYIA ALPINA hn. sp. (Figs 78-80, 102-106) 
Types: Holotype 2 and 19 paratype. 
Type Locality : Alpine Creek, Kiandra, N.S.W. (2.11.1965, D. Colless). 
A small, very dark species, the legs with narrow pale bands. Male unknown. 
Female: Length 1-44mm., wing 1:13 x 0-36 mm. 


PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, VoL. 96, Part 3 


MARGARET L. DEBENHAM 147 


Figs 78-80. Alluaudomyia alpina. 78. 2 wing. x55; 79, 2 abdomen (paratype), x 55; 
80, 2 spermathecae (paratype), 200. Figs 81-85. Alluaudomyia latipennis. 81, 2 wing 
(Hornsby specimen), « 55; 82, 2 spermathecae (Hornsby specimen), x 200; 83, ¢ hypopygium 
(Nattai R. specimen), «200; 84, g aedeagus, with apical processes twisted laterally showing 
curvature (Hornsby specimen), < 200; 85, 3 parameres (Nattai R. specimen), x 200. Figs 86-89. 
Alluaudomyia appendiculata. 86, 2 wing, x55; 87, 2 spermathecae (paratype), x 200; 88. ¢ 
hypopygium (paratype), x 200; 89, g parameres, x 200. 


PROCEEDINGS OF THE LINNEAN SocrETy OF NEw SoutH Wates, Vou. 96, Part 3 


148 THE GENUS ALLUAUDOMYIA KIEFFER 


Head dark brown. Eyes bare, contiguous, mandibular teeth 9. Palp with 
proximal segments light brown, distal segments darker, segment III with a 
small subapical pit bearing a few long sensillae (Fig. 102). Antennal segment IT 
dark brown, flagellar segments brown, IV—VII whitish basally (Fig. 103). 


Seutum dark blackish-brown, humeral areas light brown, two large spots 
near each lateral margin also paler, scutellum broadly dark brown centrally, 
paler laterally, with 4 setae, postscutellum blackish-brown, pleuron dark brown. 
Fore coxa and fore and mid trochanters light brown, mid and hind coxae and 
hind trochanter dark brown; femora and tibiae largely dark brown, the fore 
femur with base paler and a very narrow light brown preapical band, mid and 
hind femora with slightly broader pale preapical bands, fore and mid tibiae 
with light brown sub-basal and preapical bands, hind tibia with similar but 
broader bands (Fig. 104), hind tibial comb of 8 spines; hind tarsal segment I 
brown, remaining tarsi pale fuscous, apices of segments I and II and all of III, - 
TV and V darker. Claws of all legs very unequal, ratio of length of claws to fifth 
tarsal segment 16-5:8:19 in fore, 16:6:17 in mid, and 15:6:18 in hind 
(Fig. 105). 

Wing (Fig. 78) with many macrotrichia on apical half and in anal area. 
Two wing spots, one at junction of costa and R,,;, and one before r-m, veins 
slightly brownish. Haltere pale fuscous. 

Abdominal segments, except last three, white with a large, broad, T-shaped 
spot, distal three segments brown (Fig. 79). Cerci white. Spermathecae two, 
oval, subequal (Fig. 80). 

Distribution: Known only from the type locality. 

The very dark legs of this species are quite distinctive. 


12. ALLUAUDOMYIA LATIPENNIS (Skuse), comb. nov. 
(Figs 81-85, 90-95, 107-113) 
Ceratopogon latipennis Skuse, 1889, Proc. Linn. Soc. N.S.W., 4 (2nd series) : 
308. 
Didymorphleps latipennis (Skuse) Kieffer, 1906, Genera Insectorum, fase. 42: 56 ; 
1917, Ann. Mus. Nat. Hung., 15: 193. 


Bezzie latipennis (Skuse) Kieffer, 1906, Genera Insectorum, fase. 42: 193; Lee, 

1948, Proc. Linn. Soc. N.S.W., 73: 340. 

Type: Holotype 9, in the Macleay Museum, University of Sydney. All 
that remains of the type is a single mid leg (not hind leg, as stated by Lee). 

Type Locality: Berowra, N.S.W. (Masters). 

It is apparent from Skuse’s diagram and description that latipennis is a 
species of Alluaudomyia. A number of specimens in the collections of the 
S.P.H. and T.M. and the McMaster Laboratory, C.S.I.R.O. Division of Animal 
Health, agree very closely with the original description and are considered to be 
conspecific with latipennis. The species is here redescribed from a selected female 
and male collected at Hornsby, N.S.W., which is near the type locality. 

Female: Length 2-02 mm., wing 1-53 x 0-69 mm. 

Head light yellowish-brown. Eyes bare, just separated. Mandibular 
teeth 15-16. Palp slender, pale brown, segment III with a preapical sensory 
pit bearing several long sensillae (Fig. 107). Antennal segments II-III brownish- 
ochreous, segments ITV—X whitish on basal half, light brown on apical half, 
XI-XIV light brown with base whitish, XV light brown (Fig. 108). 

Scutum fuscous yellow, without pattern, scutellum fuscous yellow but brown 
centrally, with 4 strong setae, postscutellum brown, more yellowish laterally, 
pleuron fuscous yellow. All coxae fuscous yellow, trochanters fuscous; all 
femora and tibiae pale yellowish-white, femora with apex brown, hind femur 


PROCEEDINGS OF THE LINNEAN SocreTy or NEw SoutH WALES, VoL. 96, Part 3 


MARGARET L. DEBENHAM 149 


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Figs 90-95. Alluaudomyia latipennis, pupa. 
x 200; 91, profile of operculum, x 200; 


tubercles of 7th abdominal segment (Hornsby specimen), x 200; 94, tubercles of 8th abdominal 


90, g respiratory trumpet (Nattai R. specimen). 
92, dorsal tubercles of cephalothorax, 200; 93, 


segment (Hornsby specimen), x200; 95, g anal segment (Nattai R. specimen), x 200. 
Figs 96-101. Alluaudomyia appendiculata, pupa. 96, @ respiratory trumpet, x 200: 
97, operculum (paratype), x 200; 98, dorsal tubercles of cephalothorax, x 200; 99, tubercles 


of 7th abdominal segment, x 200; 100, tubercles of 8th abdominal segment, « 200; 101, ¢ anal 
segment (paratype), x 200. 


PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SoutH WaAtLgs, Vow. 96, Part 3 


150 THE GENUS ALLUAUDOMYIA KIEFFER 


also with a broad, very pale fuscous sub-basal band, all tibiae with apex brown, 
fore tibia also with a narrow sub-basal brown band and broad, very faint fuscous 
central band, mid tibia with base brown, hind tibia with a pale fuscous central 
band (Fig. 110), hind tibial comb of 8-9 spines ; tarsi whitish except for distal 
two segments on all legs, which are pale fuscous, and hind segment I, which is 
light brown. Claws of fore and mid legs unequal, those of hind legs very unequal, 
ratio of length of claws to fifth tarsal segment 34:17:27 in fore, 33:16: 27 
in mid, and 25:9: 24 in hind (Fig. 112). 

Wing (Fig. 81) with numerous macrotrichia on apical half and in anal area. 
Three wing spots, one just before r-m, one on R,, and one at the junction of the 
costa and R,,;, the latter extending slightly on to cell R;. Haltere with stalk 
white, knob fuscous. 


Abdomen yellowish-brown. Cerci white. Spermathecae two, oval, sub- 
equal (Fig. 82). 

Male: Length 1-87 mm., wing 1-27 x0-48 mm. 

Generally similar to female, differing as follows : 


Antennal plume pale brown, flagearll segments III-XII pale yellow, 
XII-XVI brown (Fig. 109). 


Legs lacking pale fuscous areas on fore tibia and hind femur, but central 
band on hind tibia darker. Claws all paired, equal and simple, slightly more 
than half the length of the fifth tarsal segment. 


Wing with fewer macrotrichia, these restricted to the anterior edge and 
apical fourth of the wing. MHaltere white. 


Abdomen light brown. Hypopygium (Figs 83-85) light yellowish-brown, 
coxites long, narrow, styles short and scarcely curved, with rather blunt apices. 
Aedeagus with a low, squared basal arch and a pointed, dorsally curved apical 
process, caudal lobe long and narrow, ventrally curved; parameres expanded 
and just touching basally, each with a detached basal arm, apex elongated into 
a narrow, tapering process, a similar process arising subapically. 


Pupa: ULight yellowish-brown, dorsum of thorax darker. Respiratory 
trumpet pale to dark brown, 0-25 mm. long in female, slightly longer in male, 
with about 45 pairs of spiracles in female and 65-75 pairs in male, arranged 
obliquely along most of its length (Fig. 90). Operculum similar to that of 
appendiculata n. sp. (see Fig. 97), median tubercle without spine, placed well 
back from distal margin, a.m. tubercles situated between lateral corners, each 
with a short, basally directed spine, rest of surface without spines, but with very 
small tubercles, these most prominent around the bases of the a.m.’s and on the 
distal margin (Fig. 91). Dorsal tubercles of cephalothorax situated as figured, 
1, 2 and 3 with very short, stout spines, 4 with a long fine seta, 5 with pore only 
(Fig. 92). Tubercles of abdominal segments 3-7 as figured, d.a.s.m. 2 and ].a.s.m. 
small with a short, stout spine, d.a.s.m. 1 with a long, fine seta, of d.p.m.’s 5 
absent, 4 and 3, when present, represented by pores only, 1 and 2 of intermediate 
size, 2 with a short spine and 1 with a long seta, l.p.m.’s 1 and 3 large, with a 
short spine, 2 small, with a long seta, v.p.m.’s of small to medium size, 1 and 3 
with a short spine, 2 with a long seta (Fig. 93). Tubercles of 8th abdominal 
segment as figured, d.a.s.m.’s and ].a.s.m. absent, as are v.p.m.’s 1 and 3, otherwise 
similar to preceding segment (Fig. 94). Anal segment with spines in a basal 
band of 2-3 rows (Fig. 95). 


Specimens examined: New South Wales: Mt. Dromedary (2 34, 9 99, 
24.xi1.1965, light trap, I. F. B. Common and M. Upton) ; Hornsby (2 99, pinned, 
11.iv.1956, 1 9, pimned, 16.1x.1956, 1 g, 1 9 both pinned, 19.ix.1956, 1 ¢, 
23.1%.1956, 1S, 28.1x.1956, 1 9, 3.x 1956; 5 OO) 8:x°1956; hieht trap, D. J. Wee; 
1 9, 9.x.1956, bred from pupa ex wet rock face, D. J. Lee and W. W. Wirth ; 


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MARGARET L. DEBENHAM 151 


Pee 9D GOL) Ga Zosel 9565 1 Gy 26-x. 1956, 1" 9, ge ne if OU Xe LOO 
light trap, D. J. Lee); Stockyard Creek, Colo Vale (1 9, 26.x.1954, ea 
hours, suction light trap, A. L. Dyce) ; Nattai eas Mittagong ais 9, 71.1964, 
light trap, D. J. Lee; 1 3g, 4.xi.1964, bred ex pupa, D. J. Lee and L. Smee ; 

1 g, 15.xi.1968, bred ex pupa, D. J. Lee and M. L. Debenham). 


Distribution: New South Wales. 


The coloration and leg banding in this species is quite variable. In the 
Hornsby specimens the thorax ranges from fuscous yellow to ochreous brown, 
in the Colo Vale specimen it is ochreous yellow, in the Nattai River specimens 
yellow with fuscous clouds and a greenish tinge, and in the Mt. Dromedary 
specimens dark, slightly ochreous brown, sometimes with a greenish tinge. 
The legs are generally almost entirely pale, often with faint fuscous markings, 
but in the Mt. Dromedary specimens the markings are quite distinct (Fig. 111). 


The paler forms of this species are readily recognizable by the almost entirely 
pale legs. The darker forms resemble appendiculata n. sp. but can be distin- 
guished by the pale bases on all femora, the absence of a distinct preapical band 
on the mid tibia, and the different genitalia. 


13. ALLUAUDOMYIA APPENDICULATA n.sp. (Figs 86-89, 96-101, 114-120) 
Types: Holotype g, allotype 9, 17 3g and 11 929 paratypes. 


Type Locality: Hornsby, N.S.W. (holotype 28.ix.1956, light trap, D. J. 
Lee, allotype 9.x.1956, bred from pupa ex wet rock face, D. J. Lee and W. W. 
Wirth). Paratypes from Hornsby (1 9, pinned, 6.ix.1956, D. J. Lee and W. W. 
Wirth; 1 4, 24.ix.1956, light trap, D. J. Lee; 1 dg, same data as holotype ; 
1 4g, 8.x.1956, light trap, Die see eles oy SD 99 (3 29 pinned), same data as 
allotype; 1 3, 10.x.1956, 1 6, 24.x. 1956, tg, f 9, 25.%.1956, 2 Sg, 26.%.1956, 
light trap, D. Jeiee 3) 1 on 22.x.1957, light ‘trap, 300 ft., D. J. Lee ; J boats ProeaGal WES 
light trap, D. J. Lee ; il 3; 2X 1957, light trap, 300 ft, Dandie Lee) and Asquith, 
N.S.W. (2 33, 2 22, 19. xi.1965, 3 33, 1 9, 22.xi1.1965, light trap, A. L. Dyce). 


A large brown and yellow species, with legs distinctly banded, and the wing 
spot at the apex of the costa extending on to cell R,. 


Male: Length 2-62 mm., wing 1-42 x0-49 mm. 


Head brown. Eyes bare, contiguous. Palp light brown, segment III 
with a shallow preapical pit bearing several long sensillae (Fig. 115). Antennal 
segment II ochreous, III—XII very pale, XIJI-XV brown (Fig. 117); plume 
brown. Al 


Scutum yellowish-brown on anterior half with a large brown spot on posterior 
half, humeral areas whitish-yellow, scutellum yellow, dark fuscous centrally, 
with 4 setae, postscutellum and pleuron brown. Fore and mid coxae yellowish, 
hind coxa and all trochanters brown ; femora and tibiae yellow, the fore femur 
fuscous basally, broadly brown centrally, and with a brown preapical ring, mid 
femur brown on basal half and at apex, hind femur brown on basal half and with 
a brown preapical ring, fore tibia with a brown sub-basal ring, broadly brown 
centrally, apex brown, mid tibia with a brown sub-basal ring, a narrow brown 
preapical band and a brown apex, hind tibia with a brown sub-basal ring, a 
brown incomplete central band and a brown apex, hind tibial comb of 8 spines ; 
tarsi whitish but apices of segments I, II and III on fore and mid legs and 
segments II and III on hind leg fuscous, basal sixth of mid segment I and all of 
hind segment I brown, segments IV and V of all legs entirely fuscous. Claws of 
all legs small, equal, just over half the length of the fifth tarsal segment, each 
with a minute external basal tooth. 

Wing with three brown spots, one just before r-m cross-vein, one on R,, 
and one at junction of costa and R,,;, the latter extending posteriorly for some 


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152 THE GENUS ALLUAUDOMYIA KIEFFER 


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MARGARET L. DEBENHAM 153 


distance into cell R,, veins streaked with brown. Macrotrichia restricted to 
the anterior edge and apex between the ends of the costa and M,,,. Haltere 
white. 

Abdominal tergites brown except for the distal edge and posterolateral 
two-thirds, which are white, anterior sternites white, posterior sternites light 
brown on anterior half, pleural membranes white. Hypopygium (Figs 88-89) 
dark brown, coxites long and slender, styles whitish, short and rather stout ; 
aedeagus deeply arched, with a long, pointed apical lobe, parameres separate, 
with a short, arcuate, detached basal arm, a long tapering subapical process 
and a broad, flattened, tapered apical extension. 


Female: Length 2:25mm., wing 1-62 x0-64 mm. 
Generally similar to male, differing as follows: 


Colouring more yellowish. Eyes just contiguous. Mandibular teeth 12 
large, 6 small. Segments I-III of palp (Fig. 114) whitish. Antennal segments 
brown, the basal ones paler, all except XV with base whitish (Fig. 116). 


Scutum more yellowish, scutellum entirely yellow. Legs (Fig. 118) banded 
asin male. Claws of fore and mid legs unequal, those of hind leg very unequal, 
ratio of length of claws to fifth tarsal segment 20: 10: 25 in fore, 27 : 13 : 29 in 
mid, 25:8: 25 in hind (Fig. 119). 

Macrotrichia of wing (Fig. 86) much more extensive, covering apical half 
and part of the anal area. Brown streaking particularly prominent at base 
of M,. 

Abdomen more yellowish. Cerci white. Spermathecae (Fig. 87) two, 
subspherical, subequal, each with a short neck. 

Pupa: Pale brown, thorax darker dorsally. Respiratory trumpet light 
to dark brown, approximately 0-26 mm. long, with 50-60 pairs of spiracles 
extending obliquely along most of its length (Fig. 96). Median tubercle of 
operculum without spine, situated well back from the distal edge, a.m. tubercles 
situated between lateral corners, each with a short, basally directed spine, rest 
of surface without spines but with very small tubercles as described for latipennis 
(Skuse) (Fig. 97). Dorsal tubercles of cephalothorax as figured, 1, 2 and 3 with 
very short, stout spines, 4 with a long, fine seta, 5 with pore only (Fig. 98). 
Tubercles of abdominal segments 3-7 as figured, d.a.s.m. 2 and l.a.s.m. small, 
with a blunt spine, d.a.s.m. 1 with a long seta, d.p.m. 5 absent, 4 with a very 
short spine, 3 with pore only, 2 with a blunt spine and 1 with a fine seta, 1.p.m.’s 
1 and 3 large, with a short, blunt spine, 2 smaller, with a long seta, b.p.m.’s 
also smaller, 1 and 3 with a short, blunt spine, 2 with a long seta (Fig. 99). 
Tubercles of 8th abdominal segment similar, but d.a.s.m.’s and l.a.s.m. absent, 
as are d.p.m.’s 3 and 4 and v.p.m. 1 (Fig. 100). Anal segment with basal band of 
spines in 2-3 rows (Fig. 101). 

Additional specimens: Queensland: Ravenshoe, nr. Tully Falls (1 9, 
4.viii.1967, light trap, R. Ellis and L. Hawkins); Mt. Crosby (1 9, in alcohol, 


Haplanation of Text figs. 102-120 


Figs 102-106. Alluaudomyia alpina. 102, 9 maxillary palp, x 200; 103, 2 antennal segments 
VIII-XV, x 200; 104, 9 femora and tibiae, x55; 105, 9 fore (right) and hind (left) tarsus V and 
claw, x200; 106, 2 genital sclerotization (paratype), x 200. Figs 107-113. Allwaudomyza 
latipennis. 107, 2 maxillary palp (Hornsby specimen), x 200; 108, 2 antennal segments 
VIII-XV (Hornsby specimen), x 200; 109, 3 antennal segments X—XV (Nattai R. specimen), 
«200; 110, 9 femora and tibiae (Hornsby specimen), x55; 111, 9 femora and tibiae (Mt. 
Dromedary specimen), «55; 112, 2 fore (left) and hind (ght) tarsus V and claw (Hornsby 
specimen), 200; 113, 2 genital sclerotization (Hornsby specimen), x 200. Figs 114-120. 
Alluaudomyia appendiculata. 114, 2 maxillary palp, x200; 115, g maxillary palp, segments 
TII-V, x 200; 116, 9 antennal segments VIII-XV, x 200; 117, 3 antennal segments XI-XV, 
x 200; 118, 2 femora and tibiae, x55; 119, 2 fore (left) and hind (right) tarsus V and claw, 
x 200; 120, 2 genital sclerotization, x 200. 


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154 THE GENUS ALLUAUDOMYIA KIEFFER 


4,.xii.1965, dusk—1.30 a.m., below dam bank amongst lantana, light trap, A. L. 
Dyce; 4 33, 2 99, all except 1 g in alcohol, 5.xii.1965, dusk-dawn, Lake 
Manchester Rd., 250 yds. from Brisbane Rd., in narrow, well-grassed, timbered 
eully, light trap, A. L. Dyce and M. D. Murray); Goodar Crossing, Weir River 
(1 g, 28.11.1953, suction light trap, 1730-2130 hours, W. HE. Poole); Noondoo 
(1 9, 16.xii.1963, light trap, A. L. Dyce and M. D. Murray); Yelarbon (3 ¢4, 
27.11.1952, 1800-2115 hours, suction light trap, W. EH. Poole). New South Wales : 
Yagobie Crossing (2 99, 3.x1.1951, suction light trap, 1920-2100 and 2315-0200 
hours, A. L. Dyce); Bundy via Moree (1 9, 20.v.1952, 1600 hours, net in creek 
bed, E. J. Reye); ‘“‘ Noonameena ’’ Station via Bingara (1 9, 24.ix.1952, swept 
in sunlight above pool, 1 9, 6.x.1952, swept from creek bank, A. L. Dyce); 
Bruxner Park, Coffs Harbour (3 99, 1.xi.1965, light trap, M. Upton); Otford 
(1 9, 5.ii1.1969), bred ex pupa from mud, M. L. Debenham, R. Russell and J. 
Citowitsch) ; Colo Vale (1 9, 17.x1.1954, suction light trap, 1930-2030 hours, 
A. L. Dyce); Nattai River, Mittagong (2 gg, 25.x.1968, D. J. Lee and M. L. 
Debenham); Minnamurra Falls (1 9, 16.xi1.1960, M. Upton). Australian 
Capital Territory : Black Mountain (1 4, 17.x.1960, light trap, I. F. B. Common) ; 
Canberra (1 3, 29.iv.1963, light trap, I. F. B. Common). 


Distribution: Queensland, New South Wales, A.C.T. 


The thoracic coloration of this species is very variable, ranging from yellow 
with very pale fuscous markings to entirely dark brown. Sometimes a greenish 
tinge is present. 

This species can be distinguished from the paler specimens of latipennis 
(Skuse) by the presence of distinct banding on the femora and tibiae, and from 
the darker specimens by the dark bases of the mid and hind femora and the 
presence of a distinct preapical brown band on the mid tibia, as well as by the 
different form of the aedeagus and the much broader apical lobe of the parameres. 
It is distinguished from the male of varia n. sp. by the absence of dark punctations 
in the humeral areas, the relatively narrow, well-defined preapical band of the 
mid tibia and the form of the genitalia. 


14. ALLUAUDOMYIA VARIA n. sp. (Figs 121-131) 
Types: Holotype 3, allotype 9 (both in A.N.I.C.), 7 gg and 1 2 paratypes. 


Type Locality: Yelarbon, Queensland (27.11.1952, suction light trap, 
1800-2115 hours, W. E. Poole). Allotype from Noondoo, Queensland 
(16.xii.1963, light trap, A. L. Dyce), paratypes from Yelarbon (5 ¢4, same data 
as holotype), Noondoo (1 3, 29-30.i11.1952, suction light trap, 2145-0130 hours, 
bored spring, W. E. Poole), Moree, New South Wales (1 9, date unknown, A. L. 
Dyce) and Mungie Bundie, Meehi R., Moree, New South Wales (1 3, 16.11.1952, 
suction light trap, 2000-2130 hours, A. L. Dyce). 

A medium-sized species, the male resembling appendiculata n. sp. in colora- 
tion and form of genitalia, the female with a distinctive wing pattern which 
readily differentiates it from other known species. The head is missing in the 
allotype. 

Male: Length 1:91 mm., wing 1-01 x0-39 mm. 

Head brown. Eyes bare, broadly contiguous. Palp whitish, segment III 
with a small preapical sensory pore bearing a few sensillae (Fig. 121). Antennal 
segment II dark brown, III—XII brown, XITI—-XV dark brown (Fig. 122) ; 
plume dark brown. 

Scutum brown, darker dorsally, humeral areas pale, dark punctations at 
setal bases, these especially conspicuous in the humeral areas, scutellum light 
brown with a broad dark band either side of centre, with 4 setae, postscutellum 
dark brown, pleuron brown. Coxae and trochanters brown, the hind pair 
darker ; legs whitish, extensively banded with brown, the fore femur broadly 


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MARGARET L. DEBENHAM 155 


Figs 121-131. Alluaudomyia varia. 121, g maxillary palp, « 350; 122, gj antennal segments 
XI-XV, x 350; 123, 2 wing, x90; 124, fg wing, x90; 125, 9 femora and tibiae, x90; 126, ¢ 
femora and tibiae, x90; 127, 9 fore (wpper) and hind (lower) tarsus V and claw, x 350; 128, © 
genital sclerotization, x 350; 129 92 spermathecae, « 350; 130, g hypopygium, =< 350; 131, 3 
paramere, x 350. 


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156 THE GENUS ALLUAUDOMYIA KIEFFER 


pale basally, with a wide brown central band, a pale preapical band and a brown 
apex, mid femur brown on basal half and at apex, hind femur with basal two- 
thirds and apex brown, fore tibia largely brown with narrow, pale sub-basal and 
apical bands, mid tibia similar but sub-basal pale band broader, hind tibia with 
base and apex brown and a brown central band (Fig. 126), hind tibial comb of 
6 spines ; tarsi pale fuscous, the distal segments slightly darker, apex of fore 
segment I, base and apex of mid segment I, and all of hind segment I, brown. 
Claws of all legs small, simple, equal, half the length of the fifth tarsal segment. 


Wing (Fig. 124) with three brown spots, one proximal to r-m cross-vein, 
one on R,, and one at junction of costa and R,,;, a8 well as a pale brown cloud 
anteriorly in cell R; beyond the end of the costa, veins slightly brownish. A 
small number of macrotrichia present on the anterior edge and apex of the wing, 
with a single row extending along the wing margin from M, to Cu,, and a few 
on the lower ends of M,, M, and M;,,. Haltere yellowish, apex of knob brown. 


Abdominal tergites whitish, each with a large, pale brown, M-shaped spot, 
sternites pale, pleural membranes pale fuscous. Hypopygium (Figs 130-131) 
dark brown, coxites long and slightly tapered, styles whitish, short, apices 
rounded and finger-like ; aedeagus with a squared basal arch surmounted by a 
pair of membranous lobes, and with a trifid caudal stem arising dorsal to the 
lobes, parameres separate, with a detached basal arm, stems broad basally, 
tapering gradually, the apex produced into a long, curving, pointed process, 
on the inner surface of which is a small, triangular flap from which arises a pale, 
short, tapered, setose process. 


Female: Length (without head) 1:75 mm., wing 1-14 0-53 mm. 
Generally similar to male, but differing as follows : 


Hind tibial comb of 8 spines. Claws of fore and mid legs unequal, those of 
hind legs very unequal, ratio of length of claws to fifth tarsal segment 20 : 9 : 24 
in fore, 22 : 10 : 22 in mid, 18: 4: 21 in hind (Fig. 127). 

Wing (Fig. 123) with three radial spots and pale cloud anteriorly in cell R; 
as in Male, but also with large spots around the wing margin, one each at the 
apices of cells R;, M,, M, and M,, and one large and several smaller along the 
margin of the anal cell, as well as a spot over the basal third of M,. Macro- 
trichia more extensive than in male, covering most of apical half of wing. 


Abdomen as in male, but distal tergites entirely brown. Cerci white. 
Spermathecae two, oval, equal (Fig. 129). 

Additional specimen: Queensland: Cunnamulla, Warrego River (1 3, 
28.11.1963, A. L. Dyce and M. D. Murray). 


Distribution: Southern Queensland, northern New South Wales. 


The association of the sexes in this species is based on the close resemblances 
in coloration and leg banding, and the presence in both sexes of the pale brown 
cloud anteriorly in wing cell R; beyond the costa. However, a much larger 
series of specimens is needed before the association can be regarded as definite. 


15. ALLUAUDOMYIA PLATIPYGA Tokunaga 


Alluaudomyia platipyga Tokunaga, 1963, Pacif. Insects, 5: 221 (g only). (Type 
locality : Maprik, Sepik District, New Guinea.) 

Characteristics: A small, dark brown species. Head brown, mouth parts 
white, eyes bare. Thorax mainly brown, lateral margins of scutum yellow, 
scutellum yellow with a dark median spot, 2 setae ; femora largely brown with 
bases ochreous and a white preapical band (very narrow in mid femur), tibiae 
largely white, bases and apices dark, fore tibia with a median dark band, mid 
and hind tibiae with a narrow dark band just before centre. Wing with three 
spots, one before r-m cross-vein and one on R, pale, one at junction of costa and 


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MARGARET L. DEBENHAM 157 


R,,5 prominent, a few macrotrichia apically in cell R;. Haltere white. Abdomen 
almost white, with very pale fuscous clouds on tergites I-IV. Hypopygium 
complex, aedeagus twice as broad as long, subsquare, with lateral caudal angles 
hooked, parameres broadly fused, each with apical part flattened, angulated and 
irregularly barbed. 


Distribution: Known only from the type locality. 
The structure of the hypopygium is unique. 


16. ALLUAUDOMYIA REYEI n. sp. (Figs 132-133, 149-154, 165-169) 
Types: Holotype 4g, allotype 9, and 15 ¢¢ and 45 9° paratypes. 


Type Locality: Darwin, Northern ‘Territory (holotype and allotype 
26—27.vii.1958, Quarantine Stn., suction light trap, E. J. Reye, paratypes 14, 
A OO, 27—28.Vi1.1956, 1 9, 1—2.x1.1957, 1 9, 2-3.xi.1957, 3 99, 15-16.xi1.1957, 4 g3, 
4 OO, 22—-23.x1.1957, 1 9, 22-23.v.1958, 1 3, 1 9, 28-29.v.1958, 1 9, 30-31.v.1958, 
1 g, 4 29, 10-11.vi.1958, 2 99, 11-12.vi.1958, 2 99, 13-14.vi.1958, 1 9, 
17-18.Vi.1958, 1 g, 4 99, 18-19.Vi.1958, 1 9, 28—-29.vi.1958, 1 9, 6—7.vii.1958, 
2 99, 9-10.Vvii.1958, 1 g, 2 99, 12-13.vii.1958, 4 3d, 6 9, same date as holotype, 
all Quarantine Stn., ight trap, E. J. Reye; 1 4, 1 9, 25—-26.xi1.1957, R.A.A.F. 
Marine Sect., E. J. Reye; 2 99, 26—27.xi.1957, Hast Arm Convent, N.J. light 
trap, KE. J. Reye; 1 3, 28-29.xii.1957, N.J. light trap, J. Dyer ; 1 9, 19-20.1.1958, 
R.A.A.F., N.J. light trap, J. Dyer). 


A medium-sized brown and yellow species, the wing with spots on the anterior 
veins only. 


Male: Length 1:56 mm., wing 0-87 x 0-32 mm. 


Head yellowish, vertex brown, clypeus light brown, proboscis yellow. 
Hyes bare, just contiguous. Palp whitish, segment V pale brown, segment III 
with a shallow preapical pit bearing a few sensillae. Antennal segment IT dark 
brown, flagellum light brown (Fig. 167); plume brown. 


Scutum ochreous with dark brown markings, the humeral areas white, 
scutellum fuscous on anterior half, whitish on posterior half, with 2 setae, post- 
scutellum brown centrally, yellowish laterally, pleuron yellowish-ochreous, with 
fuscous clouds. Coxae and trochanters pale fuscous, hind pair more yellowish ; 
femora and tibiae whitish, with brown markings, the fore femur with a broad 
brown central band and a brown apex, the mid femur with a broad, very pale 
brown sub-basal band, an equally pale brown preapical ring and a brown apex 
(Fig. 169), the hind femur with a broad brown postcentral band and a brown 
apex, the fore tibia with a brown base and apex and a broad central brown band, 
the mid tibia with a brown base and apex and very pale, narrow brown bands 
on the basal and apical thirds (Fig. 169), hind tibia very slightly fuscous basally, 
and with a brown central band and apex, hind tibial comb of 6 spines; tarsi 
whitish except hind segment I, which is brown. Claws of all legs small, equal 
and simple, just over half the length of the fifth tarsal segment. 


Wing (Fig. 133) with two brown spots, one before r-m cross-vein and one at 
junction of costa and R,,;. Only a few macrotrichia present, these restricted 
to the anterior edge of the wing. Haltere white. 


Abdomen whitish, tergites I-V with a distinct, light brown M-shaped spot 
covering most of the tergite, VI with this spot indistinct, remaining tergites 
light brown. Hypopygium (Figs 152-154) brown, coxites short, styles almost 
straight, whitish, nearly as long as the coxites ; aedeagus consisting of a simple 
arch, shallowly excavated basally, from which arises dorsally a ventrally bent, 
hook-like caudal process, parameres with long, attached basal apodemes, stems 
slender, with a pair of processes arising apically, one about twice the length of 
the other. 


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158 THE GENUS ALLUAUDOMYIA KIEFFER 


vote Y 


i vy 
‘ WV Y, 
pent PLN Y c 
<I +,/ 
Ly 


Figs 132-133. Alluaudomyia reyei. 132, 2 wing, x55; 133, gd wing, x55. Figs 134-140. 
Alluaudomyia bicornis. 134, 9 wing, x55; 135, 6 wing, x55; 136-140, pupa. 136, antero- 
median tubercle of operculum, lateral view, x 200; 137, dorsal tubercles of cephalothorax, x 200 ; 
138, tubercles of 7th abdominal segment, x 200; 139, tubercles of 8th abdominal segment, x 200 ; 
140, g anal segment, «200. Figs 141-148. Alluaudomyia fragmentum. 141, 9 wing, x55; 
142, J wing, x55; 143-148, pupa. 143, g respiratory trumpet (paratype), x 200; 144, antero- 
median tubercle of operculum, lateral view (paratype), x 200; 145, dorsal tubercles of cephalo- 
thorax (paratype), x 200; 146, tubercles of 7th abdominal segment, x 200; 147, tubercles of 
8th abdominal segment, 200; 148, g anal segment, x 200. 


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MARGARET L. DEBENHAM 159 


Female: Length 1-71 mm., wing 1-04 0-47 mm. 
Generally similar to male, differing as follows : 


Approximately 11 mandibular teeth. Eyes more broadly contiguous. 
Antennal segments III-IX with bases slightly whitish (Fig. 166). 


Coloration of thorax and legs more intense than in male, leg bands quite 
distinct, dark area of mid femur more extensive (Fig. 168); distal two tarsal 
segments slightly fuscous. Claws of all legs very unequal, ratio of length of 
claws to fifth tarsal segment 21:12:21 in fore, 21:11:20 in mid, 16:6:18 
in hind (Fig. 149). 

Wing (Fig. 132) with an extra spot, on R,. Macrotrichia extensive, covering 
the apical half of the wing and extending into the anal cell. 

Cerci white. Spermathecae two, subspherical, subequal, each with a short 
chitinized neck (Fig. 150). Genital sclerotization as figured (Fig. 151). 


Additional specimens: Queensland: Mossman Gorge (1 4, 23.iv.1967, 
D. H. Colless) ; Innisfail (1 9, Kubenangee Swamp, 13.vi.1963, H. Standfast) ; 
Magnetic Island (1 g, Horseshoe Bay, 9.vii.1952, 1100 hours, net, freshwater 
swamp, H. J. Reye); Belgian Gardens, Townsville (1 4g, 11.xi.1955, 8 gg, 12 99, 
of which 2 gg and 6 99 pinned, 13.xi.1955, 2100-2359 hours, mangrove tree, 
3 $d, 5 29, of which 2 gg and 3 99 pinned, 14.xi.1955, 1 g, 3 99, of which 1 2 
pinned, 17.xi.1955, 1920-2100 hours, flying fox bait, 13 gd, 4 99, 18.xi.1955, 
0001-0300 hours, flying fox bait, 2 gg, 1 9, same date, 0300-0500 hours, flying 
fox bait, 5 33, 3 O°, same date, 2100-2359 hours, flying fox bait, 6 33, a) EP 
20.x1. 1955, 2200-0600 hours, mango tree near polluted swamp, 2 99, 23.X1.1955, 
mango tree, all specimens light trap, A. K. O’Gower); Gilruth (2 ¢¢, 1 9, 
11.xii.1963, dusk-dawn, light trap in fowl yard, A. L. Dyce) : Yelarbon (1 9, 
27.11.1953, 1800-2115 hours, 1 9g, 27-28.111.1953, 2115-0045 hours, suction light 
trap, W. E. Poole). Western Australia: The Kimberleys Gesas: WwW. of Bedford 
Downs, 10 m. S. of Lansdowne H.S., vu.1964, light trap, R. Plumb). 


Distribution: Northern Western Australia, Northern Territory, Queensland. 


The male of this species is readily distinguishable from the males of bicornis 
and jfragmentum by the form of the genitalia, but the females of these three species 
are very difficult to differentiate. However, the form of the female genital 
selerotization is sufficiently distinctive in each of the three species to be of use in 
identification. The association of male and female in reyei is based on the 
evidence of a large series from Townsville, the males of which can all be classed 
as reyei, and the females of which all have the type of genital sclerotization 
here regarded as characteristic of the female of reyei. The sub-basal brown band 
of the mid femur also tends to be more extensive in reyet than in fragmentum 
and bicornis, but this character is difficult to use with certainty. 


17. ALLUAUDOMYIA BICORNIS n.sp. (Figs 134-140, 155-159, 170-173) 

Types: Holotype 3, allotype 9, and 4 gg and 20 $9 paratypes. 

Type Locality: Darwin, Northern Territory ((holotype 17-18.vi.1958, 
N.J. light trap, allotype, 26—27.vii.1958, suction light trap, Quarantine Stn., 
EK. J. Reye; paratypes, 1 9, 2-3.xi.1957, 1 9, 22—23.v.1958, 2 99, 27—28.v.1958, 
1 Q, 28-29.v.1958, 2 99, 10—-11.vi.1958, 2 99, 13-14.v1.1958, 1 9, 14—-15.v1.1958, 
2 99, same date as holotype, 3 gd, 4 29, 18-19.vi.1958, 1 3, 2 29, same date as 
allotype, Quarantine Stn., light trap, H. J. Reye; 1 9, 26—27.xi.1957, East Arm 
Convent, N.J. light trap, EH. J. Reye; 1 9, 28.iv.1958, N.J. light trap, J. Dyer). 

A small yellow and brown species with a two or three spotted wing. 

Male: Length 1-27mm., wing 0-81 x0-30 mm. 

Head brown. Hyes bare, just contiguous. Palp whitish, segment III 
with a preapical pit bearing sensillae (Fig. 170). Antennal segment II dark 
brown, segments IIJ—-XII whitish, segments XIII-XV brown (Fig. 172). 


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s, Vou. 96, Part 3 


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MARGARET L. DEBENHAM 161 


Seutum brown with darker brown markings, particularly around setal bases, 
humeral areas and lateral margins yellowish, scutellum brown on anterior half, 
pale on posterior half, with 2 setae, postscutellum dark brown, lighter laterally, 
pleuron brown, but yellowish immediately below scutum. Coxae and trochanters 
pale yellowish-brown ; femora and tibiae whitish, fore femur with a broad brown 
band just beyond centre and brown apex, mid femur with a pale brown, 
moderately broad central band, a narrow, incomplete preapical ring and a brown 
apex, hind femur with a moderately broad brown band just beyond centre and a 
brown apex, fore tibia with base, broad central area and apex brown, mid tibia 
with base and apex brown and a very pale fuscous preapical band, hind tibia 
with base and apex brown and a narrow, incomplete central band, hind tibial 
comb of 6 spines ; tarsi whitish, hind first tarsal segment pale fuscous. Claws of 
all legs small, equal and simple, about half the length of the fifth tarsal segment. 


Wing (Fig. 135) with two brown spots, one before r-m cross-vein and one at 
junction of costa and R,,;. Macrotrichia very sparse, restricted to the anterior 
margin of the wing apex. Haltere white. 

Abdomen pale brown, becoming slightly darker distally. Hypopygium 
(Figs 158-159) dark brown, styles white, slightly sinuous ; aedeagus with basal 
arch very shallow, scarcely excavated, apex bifid, forming a pair of horn-like 
processes, parameres with triangular bases, stems tapering, apices drawn out into 
long, tapered processes. 

Female: Length 1:29mm., wing 1-06 x0-46 mm. 

Generally similar to male, differing as follows : 

Approximately 10 mandibular teeth. Antennal segments IV—VIII with 
bases whitish (Fig. 171). 

Scutum more yellowish than in male, brown markings more distinct, humeral 
areas whitish. Leg banding more distinct, mid tibia with preapical band darker, 
also with a very pale fuscous sub-basal band, central band of hind tibia complete 
(Fig. 173), hind tibial comb of 7-8 spines. Claws of all legs very unequal, ratio 
of length of claws to fifth tarsal segment 23 : 12 : 22 in fore, 26:14 : 22 in mid, 
20:10:18 in hind (Fig. 155). 

Wing (Fig. 134) with three brown spots, one before r-m cross-vein, one on 
R, and one at junction of costa and R,,;. Macrotrichia dense, covering apical 
half of wing and extending into anal cell. 


Cerci white. Spermathecae (Fig. 156) two, subspherical, subequal, each 
with a short, stout, chitinized neck. Genital sclerotization as figured (Fig. 157). 


Pupa (Manton R. specimen): Light brown, dorsum of thorax darker. 
Respiratory trumpet shrivelled, but similar to that of fragmentum n. sp., with 
12 pairs of spiracles. Operculum with small a.m. tubercles bearing short spines 
(Fig. 136) rest not visible. Dorsal tubercles of cephalothorax as figured, 1, 2 
and 3 with a short, stout spine, 4 with a long, fine seta, 5 with pore only, 4 and 5 
more widely separated than in fumosipennis n. sp. (Fig. 137). Tubercles of 
abdominal segments 3-7 as figured, d.a.s.m. 2 and l.a.s.m. with stout, blunt spines, 
d.a.s.m. 1 with a long seta, d.p.m.’s 4 and 5 with pore only, 3 with a stout spine, 


Haplanation of Teat figs. 149-164 


Figs 149-154. Alluaudomyia reyet. 149, 2 fore (upper) and hind (lower) tarsus V and claw, 
x 350; 150, 2 spermathecae, x 350; 151, 9 genital sclerotization, x 350; 152, g hypopygium, 
x 350; 153, lateral view of aedeagal arch (Townsville specimen), x 350; 154, 3 parameres, 
x 350. Figs 155-159. Alluaudomyia bicornis. 155, 2 fore (wpper) and hind (lower) tarsus V and 
claw, <x 350; 156, 9 spermathecae, x 350; 157, 2 genital sclerotization, x 350; 158, g hypo- 
pygium (Mossman Gorge specimen), x 350; 159, g parameres (Mossman Gorge specimen), X 350. 
Figs 160-164. Alluwaudomyia fragmentum. 160, 2 fore (upper) and hind (Jower) tarsus V and claw, 
x 350; 161, 2 spermathecae, x 350; 162, 2 genital sclerotization, x 350; 163, gj hypopygium, 
x 350; 164. g parameres, x 350. 


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PROCEEDINGS OF 


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MARGARET L. DEBENHAM 163 


2 with a long seta, 1 absent, 1.p.m.’s 1 and 3 with a stout spine, 2 with a long seta, 
v.p.m.’s 1 and 3 with a spine, 2 with a seta (Fig. 138). Tubercles of abdominal 
segment 8 as figured, d.a.s.m.’s, l.a.s.m., d.p.m.’s 1 and 4—5, and v.p.m. 1 absent, 
remaining tubercles similar to those on preceding segment (Fig. 139). Anal 
segment with a narrow basal band of 2-3 rows of spines (Fig. 140). 

Additional specimens: Northern Territory: Manton R. (1 4, 6.vi.1958, 
bred ex pupa from reservoir, H. J. Reye). Queensland: Mossman Gorge 
(1 g, 23.iv.1967, D. H. Colless); Gilruth (2 gg, 15.xii.1963, light trap, A. L. 
Dyce); Yelarbon (2 99, 27.11.1953, 1800-2115 hours, 7 99, 27-28.ii1.1953, 
2115-0045 hours, suction light trap, W. EH. Poole); Noondoo (1 9, 29.11.1953, 
2145-0130 hours, suction light trap, W. E. Poole; 1 9, 26.11.1963, light trap, 
A. L. Dyce and M. D. Murray ; 1 9, 16.xi1.1963, light trap, A. L. Dyce); Texas 
Station (2 99, 26.10.1953, dusk—2110 hours and dusk—2130 hours, suction light 
trap, A. L. Dyce). New South Wales: Mungie Bundie, Meehi River, Moree 
(2 99, 16.11.1952, 2000-2130 hours, suction light trap, A. L. Dyce); Yagobie 
(1 9, 3.x1.1951, 1920-2100 hours, suction light trap, A. L. Dyce). 

Distribution: Northern Territory, Queensland, northern New South Wales. 


The male genitalia of this species are quite distinctive, but females are 
apparently distinguishable from females of reyet n. sp. and fragmentum 0. sp. 
only by the form of the genital sclerotization. The single pupa available can be 
distinguished from the pupa of fragmentum by the smaller anteromarginal 
tubercles on the operculum, the absence of the first dorsal posteromarginal 
tubercle on abdominal segments 3-7, and the lack of anterior displacement of 
the second lateral posteromarginal abdominal tubercle. 


18. ALLUAUDOMYIA FRAGMENTUM n.sp. (Figs 141-148, 160-164, 174-178) 

Types: Holotype Qg, allotype 9, and 1 g and 1 @ paratypes. 

Type Locality: Manton River, Northern Territory (holotype, 6.vi.1958, 
bred from pupa, E. J. Reye; allotypé 15.vi.1958, ex reservoir, EH. J. Reye; 
paratype ¢ 1.vi.1958, bred from pupa ex reservoir, EK. J. Reye; paratype 9 
same data as holotype, ex reservoir). 

_ A medium-sized yellow and brown species with a pale abdomen and two or 
three spotted wing. 

Male: Length 1-89 mm., wing 0-84 x 0-33 mm. 

Head brown. LHyes bare, just contiguous. Palp whitish, segments IV and V 
pale brown, segment III with a preapical pit bearing sensillae (Fig. 174). Antennal 
segment II dark brown, flagellum brown (Fig. 176); plume dark brown. 

Seutum brown with darker brown markings, particularly around setal bases, 
humeral areas and lateral margins yellowish, scutellum brown on anterior half, 
pale brown posteriorly, with 2 setae, postscutellum dark brown, paler laterally, 
pleuron brown except for a transverse yellow band just below the scutum. 
Coxae and trochanters pale brown ; femora and tibiae whitish, fore femur with 
central third and apex brown, mid tibia with a broad central brown band, a 
pale brown preapical band and a brown apex, hind femur with an almost obsolete 
pale brown central band and a brown apex, fore tibia with base, apex and broad 


EHxplanation of Teat figs. 165-178 


Figs 165-169. Alluaudomyia reyet. 165, 2 maxillary palp, x 350; 166, 2 antennal segments 
VIII-XV, x350; 167, g antennal segments XI-XV (paratype), x350; 168, 2 femora and 
tibiae, x90; 169, g mid femur and tibia, x90. Figs 170-173. Alluaudomyia bicornis. 170, $ 
maxillary palp, x 350; 171, 2 antennal segments VIII-XV, x 350; 172, g¢ antennal segments 
XI-XV, x350; 173, 9 femora and tibiae, x90. Figs 174-178. Alluaudomyia fragmentum. 
174, g maxillary palp, x 350; 175, 2 antennal segments VITI-XV, x350; 176, ¢ antennal 
segments XI-XV, x 350; 177, 2 femora and tibiae, x90; 178. g femora and tibiae, x 90. 


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164 THE GENUS ALLUAUDOMYIA KIEFFER 


central area brown, mid tibia with base and apex brown, rest very pale fuscous 
except for a whitish sub-basal and preapical band, hind tibia with base, apex and 
narrow central band brown (Fig. 178), hind tibial comb of 6 spines ; tarsi whitish 
except hind tarsal segment I light brown. Claws of all legs small, equal and 
simple, half to two-thirds the length of the fifth tarsal segment. 


Wing (Fig. 142) with two brown spots, one just before r-m cross-vein, the 
other covering R, and the apical ends of the costa and R,,;. Macrotrichia very 
sparse, restricted to the anterior margin of the apical third of the wing. Haltere 
whitish. 


Abdomen very pale, almost white, with light brown markings on tergites. 
Hypopygium (Figs 163-164) dark brown, but coxites paler on apical half, 
styles sinuous, rather stout; aedeagus with a very shallow arch, the apex 
extended into two laterally bent processes, parameres with stout stems, each 
with a short, flat apical process which terminates abruptly in a serrated edge, 
as though broken. 


Female: Length 1-78 mm., wing 1-06 x0-45 mm. 
Generally similar to male, differing as follows : 


Approximately 12 mandibular teeth. Segments IV—VIII of antennae with 
bases whitish (Fig. 175). 


Leg banding slightly more distinct, central band on hind femur not almost 
obsolete (Fig. 177), hind tibial comb of 6-7 spines. Claws of all legs very unequal, 
ratio of length of claws to fifth tarsal segment 18:7: 21 in fore, 19:8: 20 in 
mid, 16: 6:19 in hind (Fig. 160). 


Wing (Fig. 141) with three brown spots, one before r-m cross-vein, one on 
Rk, and one at junction of costa and R,,;, radial cell considerably longer than in 
male. Macrotrichia dense, covering apical half of wing and extending into 
anal cell. 


Cerci white. Spermathecae two, oval, subequal, each with a short, thick, 
chitinized neck (Fig. 161). Genital sclerotization extensive, as figured (Fig. 162). 


Pupa: Wight brown, dorsum of thorax darker. Respiratory trumpet dark 
brown, with 11-12 pairs of spiracles on apical third (Fig. 143). Median tubercle 
of operculum without spine, situated well back from distal edge, a.m. tubercles 
large, each with a short, stout spine (Fig. 144), rest of surface with small tubercles. 
Dorsal tubercles of cephalothorax as figured, 1, 2 and 3 with a short, stout spine, 
4 with a long, fine seta, 5 with pore only (Fig. 145). Tubercles of abdominal 
segments 3-7 as figured, d.a.s.m. 2 and ].a.s.m. with a short, stout spine, d.a.s.m. 1 
with a long seta, d.p.m.’s 1, 4 and 5 with pore only, 2 with a long seta and 3 with a 
stout spine, l.p.m.’s 1 and 3 with a stout spine, 2 displaced anteriorly so it is in 
line with the anterosubmarginal tubercles, with a long seta, v.p.m.’s 1 and 3 
with a stout spine, 2 with a long seta (Fig. 146). Tubercles of abdominal segment 
8 as figured, d.a.s.m.’s, l.a.s.m., d.p.m.’s 1, 4 and 5 and v.p.m. 1 absent, remaining 
tubercles similar to those on preceding segment (Fig. 147). Anal segment 
with a basal band of 2-3 rows of small spines (Fig. 148). 


Additional specimen: Northern Territory: Darwin (1 9, 22-—23.v.1958, 
Quarantine Stn., N.J. light trap, E. J. Reye). 


DMstribution: Northern Territory. 


The male of this species can be readily identified by the form of the genitalia. 
The female is apparently only distinguishable from females of reyes n. sp. and 
bicornis n. sp. by the extensive genital sclerotization. The sexes are associated 
on the basis of pupal characters, particularly the anterior displacement of the 
second lateral posteromarginal tubercle on abdominal segments 3-7. 


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MARGARET L. DEBENHAM 165 


Figs 179-184. Alluaudomyia petersi (Lae specimen). 179, 9 maxillary palp, x 350; 180, 2 
antennal segments XITI-XV. x350; 181, 9 wing, X90; 182, 2 femora and tibiae, x 90: 
183, 2 fore (left) and hind (right) tarsus V and claw, x 350; 184, spermathecae, x 350. 


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166 THE GENUS ALLUAUDOMYIA KIEFFER 


19. ALLUAUDOMYIA PETERSI Tokunaga. (Figs 179-184) 
Alluaudomyia peterst Tokunaga, 1963, Pacif. Insects, 5: 228 (2 only). (Type 
locality: Maprik, Sepik District, New Guinea; paratype from Bainyik, 

New Guinea.) 

Specimens examined: New Guinea: Damanti Village, Finisterre Range, 
Madang Central Subdistrict (1 9, x.1964, 3,500 ft., R. Pullen) ; Lae (1 9, vii.1958, 
light trap, W. Peters). Holotype also examined. 

Characteristics : A medium-sized brown and yellow species. Male unknown. 
Head ochreous, eyes bare. Thorax yellow, extensively mottled with brown, 
seutellum yellow with a median fuscous spot, 4 setae, postscutellum fuscous 
with two yellowish anterior spots; legs yellow, banding as figured (Fig. 182), 
female claws (Fig. 183) unequal, ratio of length of claws to fifth tarsal segment 
28-8:14-8: 28 in fore, 29-8:14-3: 27-5 in mid, 19-3: 8:22 in hind. Wing 
with three spots, one before r-m crosgs-vein, one on R, and one at junction of 
costa and R,,;, the latter spot extending into cell R,; (Fig. 181), macrotrichia 
numerous on apical half of wing and in anal cell. MHaltere white. Abdominal 
tergites mainly brown, but I widely yellow, II—-VIII yellow on distal margin, 
IX and caudal end white, cerci white. Spermathecae (Fig. 184) two, round, 
subequal, one with a short process, the other with a slightly longer process. 

Distribution: New Guinea. 

Tokunaga states that the macrotrichia on the wing of this species are 
‘‘rather sparsely spread on apical part beyond level of costal end ”’, and his 
diagram shows this arrangement, but a check of the holotype reveals that the 
macrotrichia are rather more extensive, agreeing with the present specimens. 
The broadly dark basal end of the mid tibia readily identifies this species. 


20. ALLUAUDOMYIA ASTERA Tokunaga 


Alluaudomyia astera Tokunaga, 1963, Pacif. Insects, 5: 222 (g only). (Type 
locality : Inis Atoll, Bougainville I., Solomon Is.) 

Characteristics: A small yellow and brown species. Kemale unknown. 
Head ochreous, eyes bare. Thorax with tergites mainly yellow, pleurites and 
sternites fuscous, scutum with a median star or crux-shaped dark spot and several 
fuscous clouds and spots, scutellum yellow with a dark median spot, 4 setae, 
postscutellum dark centrally, yellow laterally ; legs yellow with brown bands, 
fore femur with narrow pale base, median part fuscous, a narrow, indistinctly 
paler preapical ring and dark apex, tibia with base, apex and broad median part 
dark, mid femur entirely pale except for a dark preapical ring and dark apex, 
tibia with base and apex dark and a dark sub-basal and preapical band, hind 
femur widely dark medially and with a dark apex, tibia with a narrow central 
dark band and dark apex. Wing with three spots, as in peterst Tokunaga, 
macrotrichia sparsely spread on wing margin between ends of costa and M,. 
Haltere white. Abdomen yellowish ochreous, tergites I-VI with pale fuscous 
clouds, pleural membranes dark. Hypopygium yellow, coxites and tips of 
styles dark, aedeagus bell-shaped, short caudal stem bent ventrally, parameres 
broadened, flattened and oval on apical two-fifths. 


Distribution: Known only from the type locality. 


This species can be distinguished from jimmensis Tokunaga and smeei 
Tokunaga, which have similar leg banding, by the entirely pale basal two-thirds 
of the mid femur and by the more yellowish scutum. 


21. ALLUAUDOMYIA JIMMENSIS Tokunaga. (Figs 185-195) 
Alluaudomyia jimmensis Tokunaga, 1963, Pacif. Insects, 5: 229. (Type locality : 
Tsenga, Jimmi Valley, N.E. New Guinea; allotype from Enarotadi, 
Wisselmeren, West Irian, paratype from Kainantu, N.E. New Guinea.) 


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MARGARET L. DEBENHAM 167 


Specumens examined: New Guinea: Lumi (1 g, v.1969, 2,000 ft., Dr. L. 
Wark); Maprik (2 gg, 1 9, 1958). 


Characteristics : A medium-sized brown and yellow species. Head brown, 
frons yellow, eyes bare. Thorax yellow with extensive light and dark brown 
markings (Fig. 195), scutellum yellow with two fuscous spots and 4 setae, post- 
scutellum brown with two anterior yellow spots; legs yellow, with banding as 
figured (Fig. 189), female claws (Fig. 190) unequal, ratio of length of claws to fifth 
tarsal segment 28:15:30 in fore, 30:12:30 in mid, 20:?:24-5 in hind 
(20: 8-4: 24 in Maprik specimen), male claws all small and equal. Wing with 
three spots, one before r-m cross-vein, one on R,, and one at the junction of the 
costa and R,,,, the latter spot extending on to the membrane (Fig. 188), female 
with a moderate number of macrotrichia on the apical third of the wing, male 
with only a few macrotrichia on the wing margin. Haltere white. Abdomen 
pale, in female tergite I with a small fuscous cloud, II-VI with M-shaped fuscous 
fuscous markings, VII—VIII fuscous, XI and cerci white, in male tergite I fuscous 
laterally, II-III with M-shaped spot, IV—V widely dark, VI-VII fuscous laterally, 
VIII entirely fuscous, hypopygium brown. Spermathecae (Fig. 191) two, equal, 
round, one with a short, straight diverticulum, the other with a long and undulate 
process. Aedeagus triangular, apex bent ventrally, parameres separate, each 
with a detached basal arm, stem slightly swollen medially, apex flattened and 
pointed, slightly bent laterally (Figs 192-194). 


Distribution: New Guinea. 


This species is very similar to smeei Tokunaga, but the females are readily 
distinguished by the spermathecal diverticula, the longer one in jimmensis being 
undulate and about 0-05 mm. long, while in smeei it is straight and under 
0:04mm. long. The male of smeet is unknown. 


22. ALLUAUDOMYIA SMEEI Tokunaga. (Figs 196-197) 


Alluaudomyia smeei Tokunaga, 1963, Pacif. Insects, 5: 230 (Q only). (Type 
locality : Lowlands Agr. Stat., Keravat, New Britain.) 


Specimen examined: New Guinea: Maprik (1 9, iv.1958). Holotype also 
examined. 


Characteristics : A small yellow and brown mottled species. Head brown, 
eyes bare. Thorax yellow and brown, scutum yellow with an extensive brown 
pattern very similar to that in jimmensis, scutellum brown centrally, yellow 
laterally, with 4 setae, postscutellum brown with two anterior yellow spots ; 
legs yellow, banded brown as figured (Fig. 196), female claws (Fig. 197) all 
unequal, ratio of length of claws to fifth tarsal segment 20:12:19 in fore, 
20:12*:17 in mid, 14:6:16 in hind. Wing with two spots, one before r-m 
cross-vein and one at junction of costa and R,_;, the latter spot extending very 
slightly into cell R;, macrotrichia sparse, restricted almost entirely to wing 
Margin between the ends of the costa and M;,,. MHaltere white. Abdominal 
tergites white with large M-shaped brown spots similar to jimmensis. Sperma- 
theeae round, slightly unequal, in the type diverticulum of larger spermatheca, 
approximately twice the length of the diverticulum of the smaller. 


Distribution: New Britain, northern New Guinea. 


The specimen from Maprik differs in having the longer spermathecal 
diverticulum only one and a third times the length of the shorter diverticulum. 
However, it appears identical to smeei in other characters, and so at present is 
included in this species. 


* Not 19 as in original description (remeasured on holotype). 


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MARGARET L. DEBENHAM 169 


23. ALLUAUDOMYIA SPINOSIPES Tokunaga. (Figs 198-207) 

Alluaudomyia spinosipes Tokunaga, 1962, Pacif. Insects, 4: 206. (Type 
locality: Chibana, Okinawa; paratypes from Sonabi and Yaka, 
Okinawa); Wirth and Delfinado, 1964, Pacif. Insects, 6: 615 (Ceylon, 
Indonesia, Laos, Malaya, North Borneo, Philippines, Sarawak, Thailand, 
Viet Nam). 

Alluaudomyia novaguineae (also as novaguineana) Tokunaga, 1963, Pacif. Insects, 
Od: 227. (Type locality: Minj, W. Highlands, New Guinea; paratype 
from Maprik, New Guinea.) New synonymy. 

Specimens examined: New Guinea: Maprik (3 gd, 7 929, 1958; 1 9, 
iv.1958; 1 3, vu.1958). Holotype and allotype of A. novaguineae and 1 4, 
11 99, from Wirth and Delfinado’s specimens of A. spinosipes also examined. 

Wirth and Delfinado state that novaguineae (novaguineana) differs from 
spinosipes in having the female claws all unequal. An examination of the 
allotype female of novaguineae shows that the claws are, in fact, as described 
for spinosipes, i.e. fore and mid claws subequal, hind unequal. Comparison of 
descriptions and specimens of novaguineae and spinosipes reveals no differences 
that could be regarded as of specific value, so novaguineae must be considered a 
synonym of spinosipes. 

Characteristics: A medium-sized yellow and brown mottled species. Head 
brown, eyes bare. -Scutum yellow, mottled extensively with dark brown, 
scutellum yellow with a central fuscous spot, 4 setae; legs yellow with brown 
bands as figured (Fig. 201), female with fore and mid claws subequal, hind claws 
very unequal (Fig. 202), ratio of length of claws to fifth tarsal segment 20 : 20 : 23 
in fore and mid, 20:8: 21 in hind, male claws all small and equal. Wing 
(Fig. 200) with two dark spots, one before r-m cross-vein and one at junction of 
costa and R,,;, in female macrotrichia sparsely spread on anterior edge and 
apical fourth of wing, in male macrotrichia restricted to apical margin of cells 
R,and M,. MHaltere of male white, of female yellow with knob brown. Abdomen 
yellow and brown, tergite I yellow, sometimes with pale brown median spot, 
II-III yellow with brown T-shaped spot on proximal three-fourths, IV and V 
similar to III or entirely brown, VI similar to III, entirely brown, or yellow, 
VII yellow, VIII brown or yellow, [X brown or white (Fig. 203), female cerci 
white, male hypopygium brown. Spermathecae two, equal, subspherical, 
each with a short, clavate diverticulum (Fig. 204). Aedeagus bell-shaped, 
apex bent ventrally, parameres separate, stems very slender on basal half, 
broader on apical half, apices sharply bent laterally, triangular (Figs 205-207). 

Distribution: Ceylon, Thailand, Laos, Viet Nam, Ryukyu Islands, Indonesia, 
Malaya, Sarawak, North Borneo, Philippines, New Guinea. 

The subequal claws on the fore and mid legs of the female, the clavate 
diverticula of the spermathecae and the form of the male genitalia distinguish 
this species. 


24. ALLUAUDOMYIA sp. N.G. No. 1 
Alluaudomyia sp. N.G. No. 1 Tokunaga, 1963, Pacif. Insects, 5: 226 (Q only). 
(Locality : Aiyurop, 1,530 m., nr. Mendi, S. Highlands, New Guinea.) 
The leg banding and structures of the spermathecae of the single specimen 
reported by Tokunaga suggest that it could belong to an as yet undescribed 


Haplanation of Teat-figs. 185-197. 

Figs 185-195. Alluaudomyia jimmensis (Maprik specimen). 185, 2 mawnillary palp, x 350; 
186, 9 antennal segments VIII-XV, «350; 187, g antennal segments XI-XV, «350; 188, 9 
wing, X90; 189, g femora and tibiae, x90; 190, 9 fore (lower) and hind (upper) tarsus V and 
claw, ~ 350; 191, 2 spermathecae, x 350; 192, § hypopygium, x 350; 193, g paramere, x 350 ; 
194, g coxite and style, «350; 195, dorsum of thorax (Lumi specimen). Figs 196-197. 
Alluaudomyia smeei. 196, 2 femora and tibiae, x90; 197, 9 fore (right) and hind (left) tarsus V 
and claw, x 350. 


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170 THE GENUS ALLUAUDOMYIA KIEFFER 


species, but as both wings were broken off the specimen an adequate description 
cannot be made, and the species remains unnamed. 

Characteristics : A medium-sized brown species. Head brown, eyes bare. 
Thorax largely brown, scutum with indistinct paler spots, scutellum yellow, 
4 setae ; legs with all coxae, trochanters, knee parts and tibial apices dark, fore 


Figs 198-207. Alluaudomyia spinosipes (Maprik specimen). 198, 2 maxillary palp, x 350 ; 
199, 2 antennal segments VIII-XV, «350; 200, 9 wing, x90; 201, 2 femora and tibiae, x 90 ; 
202, 2 fore (left) and hind (right) tarsus V and claws, x 350; 203, 2 abdomen, x90; 204, 2 
spermathecae, x 350; 205, g hypopygium, 350; 206, g coxite and style, x350; 207, g 
parameres, xX 350. 


and hind femora fuscous on basal two-thirds, white preapically, mid femur 
fuscous on basal half, white on preapical half, tibiae largely white but fore femur 
with a fuscous median cloud, mid with a fuscous preapical cloud and hind with 
a prominent, oblique, fuscous sub-basal band, claws all unequal, ratio of lengths 


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MARGARET L. DEBENHAM 171 


of claws to fifth tarsal segment 31 : 22 : 26 in fore, 33 : 18 : 25 in mid, 21 : 11: 20 
in hind. Wing with a spot before r-m cross-vein, rest of wings broken off. 
Abdomen very pale ochreous, almost white, cerci white. Spermathecae round, 
subequal but with very unequal diverticula. 


Distribution : Known only from the locality recorded by Tokunaga. 


25. ALLUAUDOMYIA BRANDTI Tokunaga. (Figs 208-212) 


Alluaudomyia brandti Tokunaga, 1963, Pacif. Insects, 5: 223 (9 only). (Type 
locality: Kulumadau Hill, Woodlark (Murua) I., New Guinea.) 


Specimens examined: Northern Territory: Darwin (1 4, 1-2.vi.1957, 
1 9, 27—28.v.1958, 1 9, 15-16.vi.1958, 1 9, 12-13.vil.1958, Quarantine Stn., 
N.J. light trap, E. J. Reye). Holotype also examined. 


The Darwin specimens are very similar to the holotype except for the 
presence of dark dots at some of the thoracic setal bases (in the holotype only 
one or two extremely pale dots are present). However, because thoracic 
coloration is often variable within this genus, these specimens are at present 
regarded as brandiv. 


Female: Head dark brown, eyes bare. Scutum yellow, ‘anterior margin 
fuscous, humeral pits dark, W-shaped dark stripe around posterior margins of 
humeral areas, fuscous rhombic median spot, small fuscous clouds posterior to 
seutal sutures and anterior to scutellum, which is yellow, with 4 setae, post- 
scutellum brown centrally, yellow laterally ; legs yellow with brown bands as 
figured (Fig. 208), claws all very unequal (mid and hind claws missing in holotype) 
(Fig. 209). Wing with dark spots proximal to r-m and at junction of costa and 
R,,; (also a faint spot on R, in the Darwin specimens), macrotrichia spread 
over apical half of wing. Haltere white. Abdomen yellow, tergites I-VI with 
pale brown T-shaped clouds on anterior part, VIII with a dark narrow median 
band. Spermathecae two, subequal, each with a small diverticulum. 


Male: Length 1-59 mm., wing 0-84 «0-30 mm. 


Generally similar to female, differing as follows : 

Antennal plume dark brown. Thorax light brown instead of yellow, but 
with similar dark markings except some setal bases with dark dots. Claws all 
small and equal, half the length of the fifth tarsal segment. Macrotrichia 
restricted to anterior edge of wing. Hypopygium (Figs 210-212) brown, styles 
with apical third brown, sharply bent. Aedeagus bell-shaped, apical point 
bent ventrally, parameres strongly swollen on basal half of stems, apices bent 
at right angles, scarcely tapered and rather blunt. 


Distribution: Woodlark Island, Northern Territory. 


Females of brandti are distinguished from those of tenwistylata by the shorter 
spermathecal diverticula. The males are distinguished most easily by the form 
of the parameres, in brandti the stems being swollen and the apices blunt and 
scarcely tapered, in tenwistylata the stems being slender and the apices more 
Slender, tapering and undulate. 


26. ALLUAUDOMYIA TENUISTYLATA Tokunaga. (Figs 213-218) 

Alluaudomyia tenuistylata Tokunaga, 1959, Pacif. Insects, 1: 296 (g only). 

(Type locality: Hollandia [Kotabaru], West Irian); Tokunaga, 1963, 

Pacif. Insects, 5: 225 (g, 9, Kamo Valley, Itouda, Wisselmeren, West Irian 

and Maprik, Sepik District, New Guinea.) 

Specimens examined: New Guinea: Maprik (1 dg, iv.1958, 2 gg, 1958). 
Queensland: Innisfail (1 3g, Eubenangee Swamp, 14.ix.1963, H. Standfast). 
Holotype also examined. 


PROCEEDINGS OF THE LINNEAN SocreTy or NEw SoutH WaAtEs, Vou. 96, Part 3 


LA THE GENUS ALLUAUDOMYIA KIEFFER 


Figs 208-212. Alluaudomyia brandti. 208, 9 femora and tibiae, x90; 209, 2 fore tarsus V 
and claw, «350; 210, ¢ hypopygium (Darwin specimen), x 350; 211, g parameres (Darwin 
specimen). 350; 212, 3 coxite and style (Darwin specimen), 350. Figs 213-218. 
Alluaudomyia tenuistylata (Maprik specimen). 213, ¢ maxillary palp, x 350; 214, g wing, x 90; 
215, 3 femora and tibiae, x90; 216, g hypopygium, x 350; 217, g parameres, x 350; 218, 3 
eoxite and style, x 350. 


PROCEEDINGS OF THE LINNEAN SocteTy or New SoutH WALES, Vou. 96, Part 3 


MARGARET L. DRNENIAM 173 
Tani 1 : 
Meararements of Allunudomyin Specter (ma | temeciatyints Tok ! 
onnulipes oustralvenes  tokwnagoi W. and _D. Jucssipennis “unguiatyla jemarulais —iasulana Tok, and Mur. ein popuae Tok. bifaseiato Tok. latipennis (Skuse)! oppendioulata varia platings reyes tneormis Sragmentum psi oe 
g cerveunida - e c ’ 
sai 2 ? 3 e $ é 7 oF é z é 3 ¢ 3 z 3 3 é z é 2 é Z 2 é z é 2 "9 8 é z 
$ 5 
= 2 7 Bed ta Dar oo 62 2:25 OL _ LO 166 1 ba i) iso 17s 146 137 13s 205 
; ra art 1 170 teh bi 1a 1a 130 120 1 aba3 ] f See Wiican ont sgt “ 3 aera F 
Lic _ i a 033-0-35 052-0 4 
a 0 a ~ f —~ it i O-015 oo19 or O15 0.01; 0-016 0.015 0-015, oo O-ont 
eer - eg nt geen oak na OOS ME ore ey eS TP i Hy A Sk Bp 29 FER PP oR yeh eh pee cues ce et eee 
u = 00M MIDS SU LDIDIGE ses | 0tT Os aT eat TGS Midget © Solos  nineg] coloess | (prnedal colors Mutoaos Nl cufoazs ms sutuso mil (o/b | SlMmRUTOsa ia cyan mC one Oost 9-030 0k 0k 0030s SOA i Giogs = O0aT a7. 
ay a a Cf a 2 aio 0-030 te = gk ak. 
iy SEL ea eh ot a ES Ee i Me eh ey ky EH tbs Sec ere ay eet eae mT set Fel Ae He ey Ee 
Pajptint =0 bs 20 22 23 20 22 2 18 ~ = 15 = 20-23 a3 20 i) 26 25 20 20 = re) 20 25 20 18 ie rs 2 22 208 
pat rates 2 z 2 2 = 
viata srerae ee 0 = = = 0078 = = - ou 0-095 1-000 ou 0.057 = 0.083 oo4t 0.072 0.046 0.070 0-049 = ~ - = — - 0.072, = = = 
oe ie fe Eh ee as Se ee eee ey Gey ue) gat peg = See ease Cee meme c= =) — eee i a = 
on) eu = fos bond ounat = = os = = = fo onto aus Gem O00 00s oen oak one = = = = = : S00 = = = 
0) OM = bose 00 oo = = = = = 3 = E = = = 0.030 = = = 
0x0 i = = 0 bet eet = = = 030 = 030 03k 0-050 OND = = 00% = «00 = 0085 030 0-014 = = i > = - 
0.030 jos oso, mao OM) 0-027 = on = 0.02007 = CT = (020 0-05 hte 0k) 030 0038 Oras a = oss ou oma aca 0.026 = ong 4s 
pee 0026 0-023 0-020 0004 0M 0.000 0-098 0053 oo 0026 0041 0.026 0.032 035 0-042 0033, - 044 0-032 0mm 06 oom 
rea rare ici Mt Sita onan sient bent Dore yy 0-030 6-060 0-035 0053 OoLS 0090, OO 0030 0/030, 008 0-04 0-033 - 0-048 0-031 0-020, 0-030 oor - 0-030 0-055 
: i030 1:00 0:0) ke naa th08t Sis oor Osage cta on OOM ork Bote Oso = Dos bo) mdr oe baw ose Tost Oeas Oo 08 0 026 = $100 0-083 
9.030 0-030 0009 . i o-0st 0.037 mt . u ; = y ° ; : 
a comes ie sah) 8s oe Hcp eect ek Be eS eee ste ete ea a ea ated Se wats td dates > Se om 
0.072 0058 0.033 0:087 0st 00a bok 0.008 Om Ot hone OIL 078 = Dom bods sok 0-053 0-057 O08 = en oor = g-oss 9-08) 
fom hon D080 MOK 80D $060 0780-088 Oo 0106-04000 = 0-037 0-004 OAL MD HL 0:04 0-051 0-096 0 04 ome = 0-073 o-078 outs 0-084 
0:09 = ow tisk O08 i = Gat 0.0088 0se OTH = 0.08 O05; a0 BMD 10m Sa = * 
; m7 233 oa a4 
0.300 ) 3 03 oar 0285 0-318 O8IB 89a, tesa at 0.975 0-234 0315 08s 3s oe 0-38 3s 0403 a : 
0300 pass Dash 0.300 ner OHO 0263 aves SNR Oa 6300-201 O55 9.285 O3H0 O35 8-361 Ca base 0-385 pet 0-436 
o-10 . 13 ° ° 0-168 F 1m 0-404 O10 =O 108 0-208 
pe Bt pes Ane oa as eee oe sete fe soars: eg pa ae cre meee) ett re A Oe Fe 
won (0.038 om: on 010034 034 = Gos1 032.0 0038 0-00 0.03300 Oo Ob26 Goss ot fai o-ga 00a 033,039 09 8088-0928 
bom Oe 04026 8S 0026. 0-038 = f:028 b 032 034 Hi 6-08) WO 8 Dost aarp So3s 8082 a com O-nge 0038 0390-08 D038 wea gaze oun 
ban OOM 8091 ONS wR S088 = 0055083. OB (072 O02 908.088 8-095 0.030 0.070 0:03K OR rc at = 5 
re, 2 7: a 0 0-7! 0 0430 ONS 0-480, 0-300, 0-405 0-491 0-351 om 000 0-452 o-3st oon 
0 0 0486 0.003 0-300 0-510 0-480 os 04200420 85 0-404 0-600 A712 0.720 0409 ‘ 7 , 2 i 
re me oan 030 Ott OKs as jas 31s 0am Bay Bae oasis ans O43 tas a0 am am teat 031 one = ox 0am Boom 
es 227 ue 7 0 0 om ome 0193 0-205 0-195 0. 
F 0-243 om a.to1 209 oma orm a4 ga ong or 0-300 o-ai ° peas) Cony A et cals 0.008 Goss Odie O06 OTE 086 OE 
fd 0-098 0-108 0-104 0 068 0-095 0-070 0-060 0-007 O-110 0 104 0070 o- 102 0106 8s 7 7 ees 0-030 oe 0-030 0-039 0-033 0-052 
Wm Ons 0-048 os Hoa at Oa3t Cr FT 0-03R Da 6.053 6 O:038 0-045 Dos wos D0 Oos G04 a-ngvawegtggo.udy 8 
WV Diese 8:08 01030030 Om ant Ho 8.026 9032 00m 050.087 0 0-038 fms heat te 9-40 029 wosr a Gross Omak eS 
. 03k Ona Door QE 8.064 oom host 08) ge 600k O08 th 0-049 0.07 (038 Sow Gore 0a 08S 
Wend 27 2 iT 0.31 +300 0-280 a oso 0 9-300 0-364 0-264 0.507 
Waa 330 0420-480 0:50 0.300 oaxs 0390 oz ass 0338 O9BL 0387090080897 0.376 0405 0-315 0-300 2 je oe 
Tie ie eT ao 0300 842 O.a60 0-270 308982 OBR ONTOS MAR OBRSADS 0.330 0.390 SS OT OBE 0-256 as? to 4a 351 0am 0-833 
inet 3 130 O16 OTe o.20 oss Our 
163 sts 0-205 O13 O12 ote Ota ono O43 0598159205, onl 6-103 o-t48 0-108 3 ws o4 
u oss co 8076 0.038 0.004 00 5-060 0-040 0.058 0.008, 0.087 0.036 0 0.000 8-045 0004 Ovone 0-059, 048 moss) os 0-085 9-048 9-085 
Mt O00 0b 0.030 OOO b-028 00%) Ou? 88 0.038 0-026 0-004 0-00 0-030 O03 O03 Gos | 3 a8 = a3 0-088 
w O:0oK OLAS Dor? oom O03 O02 Dose 0-037 038O 0.030 6.038 0030 finxe ona D037 o.n26 9.000 0-0 = hon | Sroms 
: 00:8 0-070 Dour 0080 008k 808 0-030 0.005 O00 OOS OOO = 0-030 0-088 0.008.008 0-030 0-08 7 5 
Wier 203 1% 235 2:50 = 2:00 Las Lo 220 2.20 203 oa 2.00 to Las 2:30 os ea B 
Mid 34s 3-04 et 320 gaz 238 245 248 203 3% ao 2-01 “0 348 is 2 © Ze zs 
Hind " 233 2-60 io 24t Paty 24 2.02 280 230 2 200 i 24a 4 Re OHya ce. Ma 2s = 
a rat ob ives os O48 oa oa on 06 O40 O48 0b OW 0.98 08 2 O-8s bo Oo a ¥ vous aus cs 
Sparmatboca = = 60  o.0s30-05) 7 é = se 0-043 + 0-042 048 0.048: 0063 0-045, = 006K «D.05; —O.08RxO.0NI8, = O-DETXMONS, = * 9-041 0-019 wvosaien-os7; "=" 00485 0'1s9,0-050;60,048 00520950 0/045 0,048 "0-014 0,0414D 9 «0.04 V 0480-02 9-051 «0-051 
A +0018 0-037 x 0-026 0042 0 040; 0-049 x 0-045, “0. +0 + $0:0215 +0: . + 5 oa 
0.048 x0 041 0.040 7 0048 (0.018 0-018) 0-008 0-049 0.057 «0.047 0:05 <0: 046, 9-041 0-046 0 049 x0 053 0-041 «0-047 0-054 0-048 0.056 = 0:04 0.030 x0: 096 0-043 40,0380 51 <9. a8 0-030 0.08 0.045 0.048 
$022 0-022 0.017 70-020 40-018 2001 2 
a TMossarementa of Hornsby specimens. * Hornsby: paratype, 7 From type Of seroyuinese—not given for types of epinanipe. * From allotype of newmgvinear. Darwin specimen. 
Paoceeninos oF yuk Linxxax Soctery or New Sovrn Wates, Vou. 96, Part 3 


174 THE GENUS ALLUAUDOMYIA KIEFFER 


Characteristics: A small to medium-sized, largely yellow species. Head 
brown, eyes contiguous to just separate. Thorax almost entirely yellow, scutum 
entirely yellow or with pale fuscous clouds, sometimes with fuscous dots at setal 
bases, scutellum yellow but fuscous centrally, 4 setae, postscutellum brown 
centrally, yellow laterally, legs white to yellow, with dark bands as figured 
(Fig. 215), female claws all unequal, ratio of length of claws to fifth tarsal segment 
28:15 : 32 in fore, 29: 15 : 29 in mid, 20: 9: 26 in hind, male claws all small and 
equal. Wing (Fig. 214) with two-three spots, one before r-m cross-vein, some- 
times one on R,, and one at junction of costa and R,,, extending into cell R,, 
macrotrichia spread on apical half of wing on female, restricted to wing margin 
between the ends of the costa and M, in the male. Haltere white or with knob 
pale fuscous. Abdomen widely white, in male anterior three tergites each with 
two lateral and one median brown spots, next two tergites with a very large 
subsquare brown spot, tergite VI with a T-shaped dark spot, VII with a pair of 
lateral spots, VIII with a narrow caudal dark band, female with tergites II-VI 
and VIII with pale fuscous bands anteriorly, cerci white. Spermathecae two, 
round and subequal, each with a long, slightly curved diverticulum. Aedeagus 
(Fig. 216) bell-shaped, short caudal stem bent ventrally, parameres (Fig. 218) 
with detached basal arms, stems not swollen, apices sharply bent laterally, 
narrow, Strongly tapered, slightly undulate. 

Distribution: New Guinea, northern Queensland. 


This species can be separated from brandti Tokunaga, which has a similar 
pattern of leg banding, by the longer processes on the spermathecae and the 
more slender stems and narrower, tapering, undulate apices of the parameres. 


ACKNOWLEDGEMENTS 


J wish to thank Dr. W. W. Wirth, Systematic Entomology Laboratory, 
U.S. Department of Agriculture, and Mr. A. L. Dyce, McMaster Laboratory, 
C.8S.IL.R.0O. Division of Animal Health, Sydney, for the use of their preliminary 
notes and sketches on some of the Australian species, and to Mr. Dyce thanks 
are also due for the loan of many partially sorted specimens. The assistance 
given by Associate Professor D. J. Lee, School of Public Health and Tropical 
Medicine, Sydney, in the preparation of this paper, and the loan of type material 
from the B. P. Bishop Museum, Honolulu, is also gratefully acknowledged. 


References 
Jones, R. H., 1961.—Descriptions of pupae of thirteen North American species of Culicoides 
(Diptera: Ceratopogonidae). Ann. ent. Soc. Amer., 54 (5): 729-746. 


Lee, D. J., and Reyes, H. J., 1955.—Australasian Ceratopogonidae (Diptera, Nematocera). 
Part VII. Notes on the genera Alluaudomyia, Ceratopogon, Culicoides and Lasiohelea. PRoc. 
Linn. Soc. N.S.W., 79 (5-6): 233-246. 


SKusg, F. A. A., 1899.—Diptera of Australia. Part 6—The Chironomidae. Proc. Linn. Soc. 
N.S.W., 4 (2nd series): 286-311. 
Toxunaca, M., 1959.—New Guinea biting midges (Diptera: Ceratopogonidae). Paczf. Insects, 
1 (2-3): 177-313. 
, 1962.—Biting midges of the Ryukyu Islands (Diptera: Ceratopogonidae). Paczf. 
Insects, 4 (1): 153-217. 
, 1963.—New Guinea biting midges (Diptera: Ceratopogonidae). Pacif. Insects, 
5 (1): 211-279. 
Tokunaga, M., and Muracut, E. K., 1959.—Insects of Micronesia. Diptera: Ceratopogonidae. 
Ins. Micronesia, 12 (3): 103-434. 
WirtH, W. W., 1952.—The Heleidae of California. Univ. Calif. Pubs. Ent., 9: 95-266. 


Wirt, W. W., and Detrinapo, M. D., 1964.—Revision of the Oriental species of Alluaudomyia 
Kieffer (Diptera, Ceratopogonidae). Paczf. Insects, 6 (4): 599-648. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VoL. 96, Part 3 


SYSTEMATIC POSITION OF THE GENERA TRIODIA R.BR. AND 
PLECTRACHNE HENR. (GRAMINEAE) 


S. JACOBS 
Botany Building, School of Biological Sciences, 
University of Sydney 


[Accepted for publication 23rd June 1971] 


Synopsis 

The taxonomic position of Triodia and Plectrachne is investigated using essentially non-floral 
characters from 23 species of Triodia and two species of Plectrachne. The characters used are 
(i) morphology of the first seedling leaf, (ii) silica cells, (ii) bicellular microhairs, (iv) bundle 
sheath anatomy, (v) iodine staining, (vi) lodicules, (vii) chlorenchyma cells, (viii) starch grain 
morphology, and (ix) chromosome number and size. The usefulness and significance of these 
characters is discussed and the variation noted. From these characters it appears that the two 
genera are either Panicoid or EHragrostoid, but after consideration of floral morphology it is 
concluded that they are distinctly Hragrostoid and could perhaps be considered as primitive 
representatives of the tribe Hragrosteae. 

The validity of the generic division between the two genera is discussed and accepted from 
present data. 


INTRODUCTION 

The position of Triodia and Plectrachne within the family Gramineae has 
been a source of some disagreement. Mueller (1858-1882) implies a close 
relationship with Festuca, if only by placing some Triodia species in that genus. 
Hubbard (1934) placed them in his tribe Eragrosteae and this system was retained 
for nearly 20 years. Vickery (1953) includes Triodia amongst members of the 
tribe Eragrosteae. Gardner (1952) distributed members of the tribe Eragrosteae, 
which he did not recognize, between his tribes Festuceae and Chlorideae. He 
placed Triodia and Plectrachne in the tribe Festuceae, along with other genera 
now included in the tribe Danthoneae, and this system has been retained by 
Vickery (1961). 

These opinions have been largely based on comparative gross morphology. 
A Ithough Burbidge (1946) presented an extensive study of the leaf anatomy 
of Western Australia T’riodia species, little seems to have been done to incorporate 
these results into the classificatory system. 

Since the work of Avdulov (1931) grass taxonomy has included more and 
more microscopic characters. The incorporation of these features has solved 
some seemingly intractable systematic problems, and the present investigation 
was undertaken to determine whether these characters could clarify the systematic 
position of Triodia and Plectrachne. 


METHODS 

The references accompanying each of the methods refers to publications 

in which the basis for the descriptive technique used here is set out. 

The characters and methods used were: 

(1) The morphology of the first seedling leaves (Bor, 1960) were observed 
after germinating seeds of 7. pwngens, T. basedowti and T. irritans in 
sand. 

(2) Preparations for the study of silica cells (Metcalfe, 1960) were prepared 
by boiling portions of the leaf blade in a 1:1:1 mixture of 90 vols. 


" 


PROCEEDINGS OF THE LINNEAN Society or NEw SoutH Watss, Vou. 96, Part 3 


176 SYSTEMATIC POSITION OF GENERA TRIODIA AND PLECTRACHNE 


hydrogen peroxide: glacial acetic acid: water, until colourless, then 
transferring through a series of solutions to xylol and finally mounting 
whole in polystyrene. 

(3) Bicellular microhairs (Tateoka, 1959) which were visible in the 
preparation for the silica cells. 

(4) Bundle sheath anatomy (Brown, 1958) was determined by preparing 
transverse sections with a sledge microtome, using sections from near 
the centre of two leaves if possible ; double staining the sections with 
safranin and aniline blue, and subsequently mounting them in poly- 
styrene. 

(5) Iodine staining for location of areas of starch formation is a relatively 
new technique for which no records of any extensive investigations 
have been found to date. It was found that all sections had to be 
prepared from leaves collected immediately before staining if a consistent 
result was to be obtained. Sections were mounted in a weak iodine 
solution and no permanent mounts were prepared. 

(6) Lodicules (Tateoka, 1960) were dissected from inflorescences, freshly 
preserved in 70°% alcohol, mounted on slides, and drawn with the aid of 
a drawing apparatus. 

(7) Separated chlorenchyma cells (Jacques-Felix, 1962; Freir, 1959) were 
prepared by macerating the leaf blade in a 1:1 mixture of 10% nitric 
acid and 10°% chromic acid for approximately two to three days. After 
treatment, the material was transferred to absolute alcohol and, gently 
squashing the material, mounted in euparal. 

(8) Starch grain morphology (Tateoka, 1962) was determined by scratching 
the surface of a seed of 7. pungens and staining the starch obtained 
with a weak solution of iodine. 

(9) Chromosome number and chromosome size (Avdulov, 1931) root tips 
were treated in 8-hydroxy quinoline for 3—4 hours, macerated in a 1:1 
concentrated HCl and ethanol mixture, then stained in aceto-orcein. 

The type species for the genus Plectrachne was designated by Henrard as 

Plectrachne schinzii in Schinz. Vierteljahrs Naturf. Ges. Zurich (74: 132), whilst 
the type species for Triodia has been taken as Triodia pungens R.Br. (see 
Hitchcock, 1920). Both these species, as well as 22 other species of Triodia 
and one other species of Plectrachne, were examined for aS many of the above 
characters as possible. 


MATERIALS 


All specimens for leaf characters were obtained from the National Herbarium 
of N.S.W., the majority of which were determined by Dr. N. T. Burbidge. 

Seed material was obtained by collections made by the author for 7. basedowi 
and 7. irritans and from Dr. N. T. Burbidge for 7. pungens. 


Fresh sections for starch formation were obtained from specimens, collected 
by the author, growing in the Botany Department, University of Sydney. 


Material for the lodicules was obtained from collections made by the author 
for T. irritans and from herbarium specimens from the N.S.W. National 
Herbarium for 7’. longiceps and T. pungens. 


Material for chromosome counts was obtained from specimens growing in 
the Botany Department, University of Sydney. 


RESULTS 


(1) The morphology of the first seedling leaf has been classified into two 
main classes (Bor, 1960; Jacques-Felix, 1962) : 


PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WALES, Vou. 96, Part 3 


S. JACOBS WAH 


(i) Panicoid: characterized by an ovate to narrow-ovate first seedling 
leaf. 

(ii) Poédid: characterized by a linear first seedling leaf. 

Of these two classes, Triodia basedowti has a narrow-ovate first seedling 
leaf and is therefore Panicoid, and both Triodia pungens and Triodia irritans 
have linear first seedling leaves and are therefore Podid. 

(2) The characteristic silica cell shapes for various tribes have been described 
in detail by several authors (Duval-Jouve, 1875; Prat, 1932; Metcalfe, 1960 ; 
Bor, 1960; Jacques-Felix, 1962). In Triodia and Plectrachne several different 
forms were encountered which could be divided into two types (Fig. 1 (a)), 
viz. the basic Panicoid, or dumb-bell shape, and a series of similar types showing 
affinities with the Eragrostoid forms. Table 1 indicates an almost equal division 
of the species with Panicoid and Eragrostoid characteristics. 


S) Crs ea neiticsae en) 


C3) ROS CSRS 
(i) (ii) \ \\ 


O 
Oy aioe che wea 
0 2 9 noumie 
(iii) #4 (iv) 
, ; | 
: Wy il 
(iii) 
(ii) c 


Fig. 1. (a) Silica cells or bodies of (i) J’. concinna, (11) T. pungens, (ii) T. fitzgeraldw, and 
(iv) T. basedowti. All x 250. (i) and (ii) are characteristic Panicoid forms and (iii) and (iv) 
are characteristic Eragrostoid form. (6b) Bicellular microhairs from TJ. irritans (x 500). A 
typical Eragrostoid form. (c) Lodicules of (i) 7’. l-ngiceps (x 60), (u) 7’. pungens (x 36), (ii) 7. 
irritans (x42). Showing variations of the basic cuneate Eragrostoid form. 


The silica cells of the Plectrachne spp. examined were observed to be well 
inside the range of variation exhibited by the Triodia spp. 

Silica cell shapes are generally considered of importance as taxonomic 
characters, and presumably they are of use in the majority of cases for tribal 
classifications, but the presence of two very different forms in one genus makes 


PROCEEDINGS OF THE LINNEAN SocieTy or NEw SoutH WaAtess, Vou. 96, Part 3 


178 SYSTEMATIC POSITION OF GENERA TRIODIA AND PLECTRACHNE 


their use difficult in this particular case. On the evidence available, sub- 
family classification has two alternatives (viz. Panicoideae or Eragrostoideae), 
but Panicoid silica cell forms have been reported in sub-family Eragrostoideae 
by Tateoka (1959). 
(3) Bicellular microhairs were analysed using two ratios and a derived 

angle as an index of shape (Tateoka, 1959). The characteristics used were : 

.. Maximal width. 

@) Total length 
Length of upper cell. 
Length of lower cell | 


Maximal width —basal width x 


(il) 
1 


Length from base to rim of 
maximal width 


(iii) @=tan-!. 


TABLE | 
Grouping of Species on Silica-cell Shapes 


Panicoid-type Silica 


Eragrostoid-type Silica 


Cells Cells 

Triodia pungens Triodia basedowti 

7. mitchella hi. secunda 

a. marginata T.  fitzgeraldii 

She arritans Ts longiceps 

Oh. concinna TR. clelandir 

90. hostilis Ae wiseana 

fh. stenostachya Dy, racemigera 

Pe microstachya fs roscida 

fhe hubbardiz as mutilis 

T. spicata T.  <inaequiloba 

Ee lanata 4h. burkensis 
Plectrachne pungens Ob, intermedia 

P schinzi 


TABLE 2 
Class Limits used for the Presentation of Microhair Data (Table 3) 


Class 


Characteristics SSS Ss 
0 1 2 3 4. 


Maximal width 


poeieees ee Es 0-376—0- 500 
Total length 


0-125 0-126-0-250 0-251-0-375 0-501 


Length of upper cell 


——— a 0-800-0-301 
Length of lower cell 


1-801 1-800-1-301 1-300-1-801 0-300 


Angle between the axis of the 
hair and the line connect- 
ing the basal rim with the 


Tim of maximal width .. il" 2°—6° 7°-11° 12°-16° 17° 


Each of these characteristics was subjectively divided into classes, class 
numbers or index values being assigned to each (Table 2), and presented in the 
form A (B,C, D), where B, C, D represent index values for characteristics (1), 
(ii) and (iii) respectively, and ‘‘ A”’ represents the sum of the index values 
B, C and D (Table 3) (see Tateoka, 1959). 

The calculations for Table 2 were based on a sample of five hairs and a mean 
value taken, the variation being small. Tateoka (1959) also accepts this sampling 
method as being accurate. 


PROCEEDINGS OF THE LINNEAN Socrety or New SoutH Wates, Vou. 96, Part 3 


S. JACOBS 179 


The microhairs (Fig. 1 (b)) are all very similar, and there is a continuous 
range of variation amongst the species, but a large section of the variation of 
index numbers could be ascribed to the establishment of arbitrary classes. 


The range of values for these hairs occurs within the range of variation 
covered by the sub-family EHragrostoideae and would be classified under 
Tateoka’s tribe Chlorideae, in which he includes the two tribes Eragrosteae and 
Chlorideae of other workers. For Triodia and Plectrachne bicellular microhairs 
appear advantageous as taxonomic characters because of their relative uniformity 
between the species. It should be noted that the two species of Plectrachne 
investigated occur within the range of variation observed within Triodia. 


(4) Brown (1950) interpreted results published by Burbidge (1946) as being 
indicative of a Festucoid (Podid) leaf anatomy. A Podid leaf anatomy is 
characterized by a mestome or fibrous sheath surrounded by a parenchymatous 
Sheath, although the latter is scarcely differentiated from the surrounding 
mesophyll, and contains chloroplasts approximately the same size as those of the 
mesophyll (Bor, 1960). The drawings and photographs presented by Burbidge 
are undoubtedly accurate, but their reproduction was not intended to show this 
scale of difference and, with a more critical study of the leaf, it can be seen to 
be very different from that typified as Podid. 


TABLE 3 
Index Values from Microhair Data for the Species Examined 


Species Index Species Index 

Triodia wrritans .. 6 (2, 4, 0) Triodia concinna he (2p 2h, 1) 
T. hostilis (3) (25, 335 1) T. stenostachya TH (45 25 1) 
T. fitzgeraldi 6 (2, 3, 1) T. clelandi 7 (2, 4, 1) 
T. basedowit @ (2, Bs 1) T. racemigera UY (Bp & WW) 
T. pungens td (B & UW) T. burkensis (Bs Be 1D) 
T. longiceps 7 (3, 3, 1) T. inaequiloba 8 (3, 4, 1) 
T. intermedia 5 UW (Bs BM) T. mitchella 8 (3, 4, 1) 
T. microstachya .. 7 (B5 Bp I) T. secunda 8 (4, 3, 1) 
Plectrachne pungens t (By By 1) T. marginata SS (B, 3, 2) 
P. schinzit : WG (Ba Bo I) 


Triodia and Plectrachne are typified by relatively small areas of chlorenchyma 
enclosed by a well developed ‘‘ bundle sheath ’’ extended from around the 
vascular bundles (Fig. 2) and the relative size of the chloroplasts in the sheath 
appears to vary somewhat from being slightly larger to very much larger than 
those chloroplasts in the chlorenchyma. The range of differences between 
chloroplast sizes could be a result of drying the herbarium specimens, or of 
staining techniques. Sections of fresh material should produce more standardized 
results, but to date only fresh sections of Triodia irritans, from the Capertee 
Valley (which in fact has some characters in common with 7. scariosa, but for 
the purpose of this paper it will be referred to as 7. irritans) have been possible 
and these show a large size difference between the large sheath chloroplasts 
and the smaller ones in the chlorenchyma. This leaf anatomy could be 
characteristic of two sub-families, viz. Panicoideae and Hragrostoideae, the 
difference between the two being primarily the size difference between sheath 
chloroplasts and those of the chlorenchyma. 


When comparing differences in leaf anatomy within the genera, using the 
division as suggested and illustrated by Burbidge (1953), which is based on the 
distribution of stomatal grooves on the abaxial surface of the lamina, then three 
arbitrary classes or divisions are obtained : 


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180 SYSTEMATIC POSITION OF GENERA TRIODIA AND PLECTRACHNE 


(i) Stomatal grooves present all over the abaxial surface of the lamineae. 
(ii) Stomatal grooves present only below the midrib on the abaxial surface. 
(iii) Stomatal grooves absent on the abaxial surface. 


Class (i) (Table 4) can be further subdivided on the relative position of the 
vascular bundle as most of this group have bundles in the centre (T.S.) of the 
blade, but two, viz. 7. fitegeraldii and T. molesta, have bundles much nearer to 
the adaxial surface (Fig. 2 (b)). As well as these, there is one species, 7. 
racemigera, whose bundles are slightly closer to the adaxial surface, i.e. it forms 
an intermediate form between 7. fitegeraldui, T. molesta and the remainder of 
the class. It is suggested that the bundles being closer to the adaxial surface 
could have been the first change leading to the loss of chlorenchyma (and 


b(i) c(i) 


; Fig. 2. (a) T. hostilis (x 140), (b) (i) 7. fitzgeraldii ( x 138), (ii) T. fitzgeraldii ( x 80), (ce) (i) 7. 
irritans (111), (11) 1. irritans (200). For all figures e=chlorenchyma, v=vascular bundle, 
S=parenchymatous “‘ bundle”’ sheath. The completely inked-in areas are sclerenchyma. 


stomata) on the lower surface, which would be an obvious ecological advantage 
for leaves capable of becoming terete under dry conditions. Using this 
hypothesis, a series of changes could be drawn up, culminating in the 7. hostilis 
leaf form, perhaps representing an evolution of the leaf anatomy in response to 
the arid environment of the Australian interior. The TZ. hostilis leaf form is, 


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S. JACOBS 181 


in fact, very similar to the 7. pungens leaf form and different specimens of 
T. microstachya have both types. In the 7. hostilis leaf (Fig. 2 (a)) the “‘ bundle 
sheath” seems to be highly developed and completely surrounding the 
chlorenchyma below the stomatal groove, with only a small portion of the 
sheath coming into actual contact with the bundle. The validity of even 
considering this sheath as a ‘‘ bundle sheath” is discussed later. 


Between the two genera Triodia and Plectrachne there is a range of anatomical 
forms with no particularly distinctive character separating the two genera. 

(5) The utilization of starch formation detected by iodine staining as a 
taxonomic character has been mentioned in the literature (Brown, 1958; Bor, 
1960) but has not assumed any importance. There are two main classes of 
reaction to the staining of fresh leaves with iodine : 


(i) Members of the Podideae stain throughout the chlorenchyma including 
the bundle sheath, whilst 


(ii) Members of the Panicoideae and Eragrostoideae only stain in the 
parenchymatous sheath. 


TABLE 4 


Division of Species on the Distribution of Stomatal Grooves. Class (t) 

Stomatal grooves present all over the abaxial surface of the laminae. Class (1) 

Stomatal grooves present only below the midrib on the abaxial surface. 
Class (wi) Stomatal grooves absent on abaxial surface 


Class (i) Class (11) Class (iii) 
T. concinna T. hubbardi T. hostilis 
T. inaequiloba T. stenostachya T’. microstachya 
T. secunda T. marginata 
T. clelandia T. pungens 
T. longiceps T. mitchelli 
T. wiseana P. schinzw 
T. wrritans P. pungens 
T. burkensis 
T. lanata 
T. spicata 
T. roscida 
T. intermedia 
T. basedoww 
T. racemigera 
T. inutilis 


*T’. molesta 
*T. fitzgeraldv 


* Bundles towards adaxial surtace. 


As well as these classes Phragmites sp. (Arundineae) stains only in the 
chlorenchyma with no staining in the sheath. The differential reactions to 
iodine staining appear to be related to the C, and C, photosynthetic pathways. 


Triodia irritans (from Capertee Valley) was obtained, sectioned, and stained, 
revealing starch formation as being limited to the bundle sheath under high light 
intensities, indicating Panicoid or Eragrostoid affinities. For comparison, 
transverse sections of the leaf blade of a species of Danthonia were cut and 
stained. These showed Podid affinities by staining throughout the chlorenchyma 
and parenchymatous sheath under high light intensities. 


(6) Tateoka (1960) discusses the use of lodicules as a taxonomic character, 
but he only gives descriptions of a few tribes, a notable exception being tribes 
of the sub-family Panicoideae. From Tateoka’s descriptions, the lodicules of 
Triodia (Fig. 1 (c)) would be characteristic of members of the tribe Eragrosteae, 
but only in comparison with other members of the sub-family Eragrostoideae 
and Podideae. Prior to the use of lodicule morphology, number of lodicules 


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182 SYSTEMATIC POSITION OF GENERA TRIODIA AND PLECTRACHNE 


had been used to a limited extent, but only for a few tribes. Triodia has two 
cuneate, fleshy lodicules with fairly well developed vascularization. 


(7) The value of the shape of the chlorenchyma cell as a taxonomic character 
has been elucidated by Freir (1959) and Jacques-Felix (1962). Four characteristic 
forms have been described by these authors : 


(i) Poédid (Festucoid). Typical parenchymatous cells being approximately 
isodiametrie with joining lobes. 
(ii) Bambusoid-oryzoid. Lobed-tabular cells. 


(iii) Panicoid. Ratchet cells with deep constrictions and large angular 
axes arranged radially. 


(iv) Arundinoid. Ratchet cells arranged diffusely. 


Of the forms described above, the chlorenchyma cells of Triodia would be 
classified as Podid, although attention must be drawn to the fact that typical 
forms of the sub-family Eragrostoideae have hardly been investigated. One 
species of Hragrostis (aff. brownii) was examined and found to have Podid 
chlorenchyma cells. Further elucidation of chlorenchyma cell morphology as a 
taxonomic character is essential prior to its widespread adoption. 


(8) Tateoka (1962) re-evaluated the divisions of grass tribes based on starch 
erain morphology and concluded that simple, rounded, starch grains could be 
separated from compound, or simple, angular starch grains, resulting in two 
major divisions. The former grain type is characteristic of the sub-family 
Podideae, whereas the remaining forms seem to occur throughout the non-Podéid 
tribes. The starch grains of 7. pungens, T. irritans and T. basedowti are all 
simple and angular, i.e. they are non-Podid starch grains. 


(9) Accurate chromosome counts were difficult because of the unsuitable 
anther material collected and the thickness of the root tips to be examined. 
Associated with this was the relatively small number of cells actually dividing. 
The chromosome size was very small (i.e. non-Podid—Bor, 1960) and this also 
complicated accurate counting especially with the rather large numbers found. 
The numbers quoted can only be given as approximate because of the difficulties 
mentioned ; Triodia basedowit had 2n=60, Triodia irritans had 2n=30. As 
can be seen with such numbers, arguments based on tribal characteristics of 
n=5, n=6, n=7, n=9 or n=10 tend to be relatively meaningless as by various 
hypotheses of chromosomal losses or gains it is possible to suggest any of these 
base numbers as being the original base number before polyploidy. 


DISCUSSION 

Of the characters investigated, those generally considered to be of greater 
taxonomic significance are silica cells, bicellular microhairs, leaf anatomy and 
chromosome number. The results from the silica cells, leaf anatomy and 
chromosome size suggest Panicoid or Eragrostoid affinities, whereas the bicellular 
microhairs are more specific, indicating Eragrostoid affinities. The large 
variation of silica cell types within those genera increases the difficulty with 
which it can be applied satisfactorily, and it is suggested that silica cell shape 
can only satisfactorily be applied in conjunction with numerous other characters, 
and any attempt at identification purely by this character would be unsatisfactory. 

Bicellular microhairs appear to be reasonably characteristic even to tribal 
level and, because of their uniformity, appear to be of value for application to 
this particular problem. The microhairs are obviously Hragrostoid and, by 
comparison with the sketches and data of Tateoka (1959), are more characteristic 
of the tribe Eragrosteae, as opposed to the tribe Chlorideae. 

The anatomy of the leaf, with particular reference to bundle sheath anatomy, 
has been considered of importance since the data published by Avdulov (1931). 


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S. JACOBS 183 


From this period the concept of a bundle sheath has never been questioned, 
but evidence from Triodia and Plectrachne (also Festuca littoralis) indicates 
that the parenchymatous sheath is more closely associated with the chlorenchyma 
and its function rather than the vascular bundle. In the majority of grasses 
the chlorenchyma and the vascular strands are separated by a parenchymatous 
sheath of some form and, since it surrounds the bundle, it was considered as a 
bundle sheath. Changing the concept from a bundle sheath to a chlorenchyma 
sheath has no effect on the anatomical characters used in taxonomy, but it will 
have important implications of physiological characters, e.g. that of starch 
formation and the associated biochemical pathways. The importance of size 
differences and functional differences between sheath chloroplasts and those of 
the chlorenchyma could perhaps be better appreciated by regarding the sheath 
as being more closely associated with the chlorenchyma. 

The structure of the chlorenchyma, whether radially or diffusely arranged, 
has been utilized in conjunction with sheath type, but with 7riodia the inter- 
pretation is difficult due to its remoteness from the vascular bundle. 

Evidence from studies on the first seedling leaf has little reliability. The 
number of lodicules has been utilized in the delimitation of some groups (e.g. 
Bambuseae, Stipeae), but the variation is not sufficient for critical analysis to be 
applied. More critical studies are possible with lodicule morphology and the two 
lodicules in T'riodia show distinct Eragrostoid affinities. 


TABLE 5 
Summary of Results of Characters Investigated 


Character Danthonia Eragrosteae -Paniceae Poaeae Triodia- 
Plectrachne 
Silica cells .. a .. Panicoid KEragrostoid Panicoid Pooid Eragrostoid 
+ Panicoid 
Bicellular microhairs .. Panicoid Eragrostoid Panicoid Pooid Eragrostoid 
Bundle sheath anatomy .. Pooid Eragrostoid -Panicoid Pooid Eragrostoid/ 
Panicoid 
Lodicule shape 54 .. Oblong Cuneate — Two-lobed Cuneate 
Danthoneae Eragrostoid Pooid Eragrostoid 
Chlorenchyma cell shape .. Pooid Pooid Panicoid Pooid Pooid 
Iodine staiming gs at Pooid Panicoid Panicoid Pooid Panicoid 
Starch grain morphology .. Pooid Panicoid Panicoid Pooid Panicoid 


Morphology of first seedling 


leaf Pooid Pooid Panicoid Pooid Pooid 


The use of chlorenchyma cell shape is a relatively new technique, but suffers 
the disadvantage of having an almost continual gradation between forms, 
rendering them of limited value in many cases. Starch grain type was considered 
of prime importance prior to the work of Tateoka (1962), who has since cast 
doubt on their assumed importance. 

Staining fresh leaves with iodine is one of the simpler physiological 
characters which can be readily tested, and could become increasingly important 
in grass systematics once sufficient information has been obtained on the subject. 

Considering all the characters (Table 5) investigated, the most satisfactory 
course appears to be to include Triodia in sub-family Eragrostoideae or 
Panicoideae. To this point only leaf characteristics have been mentioned 


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184 SYSTEMATIC POSITION OF GENERA TRIODIA AND PLECTRACHNE 


(except for lodicules and chromosomes), but spikelet characteristics surely deserve 
consideration. On casual observation spikelet morphology indicates no apparent 
relationship to the Panicoideae at all, but, in some species, notably 7. irritans 
(Capertee Valley), there is an apparently classical Eragrostoid type spikelet 
structure with numerous fertile florets with the lemma three-nerved. The 
lemmas of Triodia are often apparently three-nerved, but this nervation consists 
of three groups of three traces with the laterals (of each group) being lost in 
some cases (Burbidge, 1953), as in the Triodia from the Capertee Valley. It is 
suggested that this could represent a primitive condition in the sub-family 
Eragrostoideae (in lemma characters only) supporting Hartley’s (1960) estimation 
of the centre of origin of the sub-family Eragrostoideae as being the arid regions 
of either Australia or South Africa. 


CONCLUSIONS 
Sub-family classification would place Triodia and Plectrachne in the sub- 
family Eragrostoideae, but tribal classification is more difficult. It is suggested 
that rather than attempting to establish new tribes, these genera should be placed 
in the tribe Hragrosteae, perhaps representing a ‘‘ primitive ’’ condition within 
that tribe. 


Burbidge (1953) expresses the view that T'riodia should be placed in the 
tribe Danthoneae which, in Australia at least, appears somewhat heterogeneous 
but microhair characteristics, as well as silica cell shape, iodine-staining results, 
leaf anatomy, etc., prevent it (Danthoneae) from being included in the sub-family 
Eragrostoideae. Arguments based on glume lengths and form of lobing appears 
rather arbitrary due to the large variation of these characters within the genus 
Triodia. Furthermore, the chromosome numbers reported are sufficiently 
large to accommodate any decisions based on base number (Eragrosteae : 
n=10 (5?)12; Danthoneae: n=6, 9, 12—Bor, 1960). 

Another problem worthy of consideration is the validity of the generic 
division between T'riodia and Plectrachne. The establishment of a genus is 
based on a set of correlated characters which, initially, were concerned mainly 
with the inflorescence, and, using only these characters, a generic division is 
justifiable. However, it can be seen that there are no apparent discontinuities 
for the sets of characters investigated and, indeed, the characteristics of 
Plectrachne spp. are usually well within the range of variation exhibited by 
Triodia. Taxonomically, the two genera are very close, but it is suggested that 
the generic division be maintained even if only in consideration of the large 
number of species. This division could only be maintained if no intermediate 
lemma form is discovered, but, if a lemma is found with long lobes tapering into 
small awns with a comparatively small, slightly indurated base, then the generic 
division would be invalidated since this is the sole feature which distinguishes 
the two genera. 


ACKNOWLEDGEMENTS 
I would like to express my sincere thanks to Associate Professor R. C. 
Carolin for help in carrying out these studies and help in the presentation of 
these data. 


Avputoy, N. P., 1931.—Karyo-systematische untersuchung der familie gramineen. Bull. Appl. 
Bot. Suppl., 43. 

Bor, N., 1960.—The Grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, London. 

Brown, W. V., 1950.—A cytological study of some Texas gramineae. Bull. Torrey Bot. Club, 
78: 292-299. 

, 1958.—Leaf anatomy in grass systematics. Bot. Gaz., 119: 170-178. 


- Burpipes, N. T., 1946.—Morphology and anatomy of the Western Australian species of T’riodia 
R.Br. If. Internal anatomy of leaves. Trans. Roy. Soc. S. Aust., 70 (2): 221-224. 


, 1953.—The genus Triodia R.Br. (gramineae). Aust. J. Bot., 1: 121-184. 


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S. JACOBS 185 


Duvat-Jouve, 1875.—Histotaxie des feuilles de graminees. Am. Sci. Nat., B.Ser. 6,1: 294-371. 
Freirn, F., 1959.—Las cellulas clorenquimaticas del mesofolio de las gramineae. Revista 
Argentina de Agronomia, 26: 1-16. 


GARDNER, C. A., 1952.—Flora of Western Australia, Vol. I, Part I, Gramineae. Govt. Printer, 
Perth. 


Hartiey, W., and SiatTerR, C., 1960.—Studies on the origin, evolution and distribution of the 
gramineae. III. The tribes of the sub-family eragrostoideae. Aust. J. Bot., 8: 256-276. 


Hitcucock, A. S., 1920.—U.S.D.A. Bulletin, no. 772, p. 74. 
HuBBARD, C. H., 1934.—Gramineae. In Hutchinson, Families of Flowering Plants. London. 


JAcQuES-FELIx, H., 1962.—Les Gramineaes D’ Afrique Tropicale. I. Generalites, Classification, 
Description des Genres. O.R.S.T.O.M., Paris. 


Mercatre, C. R., 1960.—Anatomy of the Monocotyledons. I. Gramineae. Oxford Press. 
Muetter, F. 8. H. von, 1858-1882.—Fragmenta Phytographieae Australiae, 8: 127-129. 


Prat, H., 1932.—L’Epidermis des gramineaes. Htudes anatomique et systematique. Ann. 
Sci. nat. Bot., Ser. 10, 14: 117-324. 


TatTEoKA, T., 1960.—Notes on some grasses. X. Some thoughts on festuceae, festucineae 
with special reference to their morphology. Can. J. Bot., 38: 911-967. 


, 1962.—Starch grains of endosperm in grass systematics. Bot. Mag. Tokyo, 75: 
377-383. 


TaTEoKA, T., INouE, S8., and Kawano, S8., 1959.—Notes on some grasses. IX. Systematic 
significance of bicellular microhairs of leaf epidermis. Bot. Gaz., 121: 80-91. 


VickEeRY, J. W., 1953.—A census of the native gramineae of New South Wales. Contr. N.S.W. 
Nat. Herb., 2 (1): 67-87. 


, 1961.—Gramineae. Contr. N.S.W. Nat. Herb., Flora Series No. 19, Part 1. 


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APHIDS OF PAPUA AND NEW GUINEA: AN ANNOTATED 
CHECK LIST 


K. P. LAMB 


Department of Biology, University of Papua and New Guinea, 
Port Moresby, T.P.N.G. 


[Accepted for publication 21st July 1971] 


Synopsis 
Fifty-seven species of aphids are listed from New Guinea. Most of these have been part of a 
collection of 6,000 alates from Moericke traps located in four different areas. Some host informa- 
tion is given, together with an index of host plants. The British Museum (Natural History) 
collection has been examined. 


INTRODUCTION 


During the past three years, more than 6,000 specimens of alate aphids 
have been collected at four main sites in the Territory of Papua and New Guinea. 
Since no previous lists of aphids have been published for this country, the 
systematic results of trapping are summarized here, together with limited host 
information, mainly based on materials in the collection of the British Museum 
(Natural History). Some records from earlier literature are included, but species 
not seen by the author are marked with an asterisk (*). Fifty-five species are 
listed. Sixteen of these species were mentioned by Eastop (1966) as occurring 
in Papua or New Guinea. 


Trapping was carried out using yellow Moericke trays at Waigani (near 
Port Moresby), Central District ; at Bulolo (altitude 2,400 ft.), Morobe District ; 
at Goroka (altitude 5,300 ft.), Eastern Highlands District; and at Keravat 
(altitude 50 ft.), Hast New Britain District. Trapped species are marked “ T ” 
and specimens examined in the British Museum Collection are marked ‘‘ BM ”’. 


Family APHIDIDAE 
Subfamily Aphidinae 
1. Aphis (Pergandeida) craccivora Koch, 1854. 

BM; T—Waigani, Bulolo, Goroka, Keravat, also specimens from 
Morobe and Aiyura. Hosts: Vigna sinensis sesquipedalis (yard-long bean), 
Vigna sinensis sinensis, Phaseolus sp., Centrosema pubescens, Gliricidia 
sepium, peanut. Hastop (1966). 

2. Aphis (Cerosipha) glycines Matsumura, 1917. 
BM; T—Waigani, Bulolo, Keravat. Host: Soya bean. 
3. Aphis (Cerosipha) gossypii Glover, 1877. 

BM; T—Waigani, Goroka, Bulolo, also specimens from Morobe and 
Chimbu. Hosts: Carica papaya, Cassia sp., Albizia chinensis, Cucurbita 
moschata, Hibiscus abelmoschus, Hibiscus rosa-sinensis, Cyphomandra (tree 
tomato), potato, taro, Tephrosia sp. Heie (1967), Valoka, New Britain ; 
Szent-Ivany (1956) on Malva parviflora. One of the most abundant and 
widespread aphid species. 

4, Aphis ? hardyi Hastop, 1966. 

T—Bulolo. 


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14. 


15. 


16. 


UG 


18. 


Wg) 


20. 


Zils 


22. 


23. 


24. 


25. 


kK. P. LAMB 187 


. Aphis nerti Boyer de Fonscolombe, 1841. 


BM; T—Waigani, Bulolo, Heie (1967)—Asclepias curassavica L. 
Lorengau, Manus, Admiralty Is. 24.vi.1962. 


. Aphis (Medoralis) spiraecola Patch, 1914. 


BM; T—Waigani, Bulolo, Keravat. Hastop (1966). 


. Brachycaudus helichrysi (Kaltenbach, 1843). 


BM; T—Waigani, Goroka, Keravat; also recorded from Aiyura. 
Host: Crotalaria anagyroides. Szent-Ivany (1956). 


. Brachysiphoniella montana (van der Goot, 1917). 


BM; Aiyura, on Leersia hexandra. 


. Capitophorus hippophaes mitegoni Hastop, 1956. 


BM; T—Waigani. 


. Hyalopterus pruni (Geoffrey, 1762). 


BM; T—Waigani, Bulolo. Host: Celosia argentea (banana). Eastop 
(1966). 


. Hyperomyzus carduellinus (Theobald, 1915). 


T—Bulolo. 


b] 


. Hysteroneura setariae (Thomas, 1877). 


BM; T—Waigani, Bulolo; also specimens from Gaimazung. 


. *Indomegoura indica (van der Goot, 1916). 


Calilung (1967) refers to this species as occurring in “ Dutch New 
Guinea ”’. 
Lipaphis erysimi (Kaltenbach, 1843). 
BM; T—Waigani, Bulolo, Keravat. 
Longiunguis sacchari (Zehntner, 1897). 
BM; T—Waigani, Keravat ; also specimens from Bulia on sugar-cane. 
Hastop (1966). : 
Macromyzus polypodicola (Takahashi, 1921). 
B T—Waigani, Goroka, Bulolo, Keravat. 
Macrosiphoniella (Pyrethromyzus) sanborni (Gillette, 1908). 
BM; Aiyura, on Chrysanthemum sp. Eastop (1966). 
Macrosiphum (Sitobion) graminis Takahashi, 1950. 
BM ; T—Goroka; Mt. Kaindi, on grass (B. C. Peters). 
* Macrosiphum (Sitobion) leelamaniae Kan. David, 1958. 
Bulolo, on grass (B. C. Peters). 
Macrosiphum (Sitobion) ? rosaeformis Das, 1918. 
T—Goroka. 
* Macrosiphum (Sitoion) ? smilacifoliae Takahashi, 1921. 
Recorded by Heie (1967) from Yalom, New Britain. 
Macrosiphum rosae (l., 1758). 
BM; Kainantu and Aiyura, on rose. 
Micromyzus dispersum (van der Goot, 1917). 
(Syn. M. eastopi Carver, 1965). 
BM; T—Goroka. 
Micromyzus ? katoi (Takahashi, 1925). 
On Platycerium wandae (syn. P. wilhelminae-reginae), Bulolo (B. C. 
Peters). 


Myzus ornatus Laing, 1932. 
BM; Aiyura, on Crotalaria anagyroides. 


? 


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188 


3T. 


38. 


APHIDS OF PAPUA AND NEW GUINEA 


. Myzus (Nectarosiphon) persicae (Sulzer, 1776). 


BM ; T—Bulolo, Goroka, Keravat ; also recorded from Aiyura. Hosts: 
Crotalaria anagyroides, Cyphomandra, Gladiolus, Ipomoea batatas, Passiflora 
foetida, Vitis arborea (pepper vine). EHastop (1966). 


. Neomyzus circumflexus (Buckton, 1876). 


BM; Aiyura, Wau, Bulolo. Hosts: Physalis peruviana, Smilax sp. 
EKastop (1966). 


. Pentalonia gavarri Kastop, 1967. 


BM; T—Keravat. 


. Pentalonia nigronervosa Coquerel, 1859. 


BM; T—Waigani, Bulolo, Goroka, Keravat. Hosts: Banana, Colo- 
casia, Caladium. 


. Rhopalosiphum maidis (Fitch, 1856). 


BM; T—Waigani, Bulolo, Keravat; also recorded from Asaro Valley, 
Aiyura, Tapini. Hosts: Sorghum vulgare, Zea mays. Szent-Ivany (1958). 


. Rhopalosiphum rufiabdominalis (Sasaki, 1899). 


T—Bulolo. 


. Schizaphis cyperi (van der Goot, 1917). 
B 


M; T—Bulolo. 


. Schizaphis graminum (Rondani, 1852). 


T—Keravat. 


. Schizaphis minuta (van der Goot, 1917). 


BM; T—Waigani. 


. Shinjia pteridifoliae (Shinji, 1929). 


T—Goroka. 


M; 


. Sinomegoura citricola (van der Goot, 1917). 


BM; Kainantu, on Avocado. Eastop (1966). 
Toxoptera aurantw (Boyer de Fonscolombe, 1841). 
BM; T—Waigani, Bulolo, Keravat; also recorded from Aiyura, 
Bubia, Goroka, Madang, Morobe, Popondetta. Hosts: Amaranthus, 
Cacao, Coffea arabica, pepper vine, tea. Dun (1954). 
Toxoptera citricidus (Kirkaldy, 1907). 

BM; T—Waigani; also recorded from Aiyura, Asaro, Bubia, Rigo and 
Wau on Citrus. EHastop (1966). 


Subfamily Lachninae 


39. 


40. 


41. 


42. 


Lachnus tropicalis (van der Goot, 1916). 
BM; Goroka on Casuarina equisetifolia. 
Greenidia ficicola Takahashi, 1921. 
BM; Eastern Highlands, on Ficus. Eastop (1966). 
Greenidia (Trichosiphum) formosana (Maki, 1917). 
BM; T—Bulolo, Wau. 
*Paratrichosiphum ? javanicum Ray Chaudhuri, 1956. 
T—Goroka, Wau. 


Subfamily Greenideimae 


43. 


44, 


45. 


Anomalosiphum pithecolobi Takahashi, 1934. 
M; T—Waigani. 
Cervaphis rappardi H. BR. Lambers, 1956. 
BM; Keravat, on cacao flowers. 
Schoutedenia lutea (van der Goot, 1917). 
BM; Port Moresby, on Breynia cernua. 


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k. P. LAMB 189 


Subfamily Drepanosiphinae 
46. Neophyllaphis araucariae Takahashi, 1937. 
BM; T—Bulolo, Goroka; Bulolo, on Araucaria cunninghamii (B. C. 
Peters); Wau. 


47. *Neophyllaphis rappardi Hille Ris Lambers, 1967. 
On Agathis labillardiert, Bosniak, Biak, Western New Guinea (Hille 
Ris Lambers, 1967). 


48. *Sensoriaphis sp. nov. 
On Nothofagus sp., Mt. Kaindi (T. Tigner). 


Subfamily Thelaxinae 
49. Kurisakia onigurumi (Shinji, 1923). 
T—Wan (Y. Lubin). 


Subfamily Hormaphidinae 
50. Astegopteryx nipae (van der Goot, 1917). 
T—Goroka. 
51. Astegopteryx rappardi H. R. Lambers, 1953. 
BM; Morobe, on coconut. Szent-Ivany (1961). 
52. Ceratovacuna lanigera Zehntner, 1897. 

BM; T—Waigani, Keravat; also recorded from Aiyura, Manum, 
Samarai, on Saccharum robustum (sugar-cane). Szent-Ivany and Cattley 
(1966). 

53. Cerataphis variabilis H. R. Lambers, 1953. 

BM; T—Keravat; Kokopo, Popondetta, Wewak. Hosts: Coconut, 
cacao. Szent-Ivany (1961). 

54. Pseudoregma bambusicola (Takahashi, 1921). 

BM; Southern Highlands, on bamboo. 


Subfamily Pemphiginae 
55. Tetraneura radicicola Strand, 1929. 
BM; Kainantu, on roots of Imperata. Szent-Ivany (1956). 


56. Tetraneura ngriabdominalis (Sasaki, 1899). 

BM; T—Waigani, Bulolo, Goroka, Keravat; also recorded from 
Galmazug and Kainantu. Hosts: grass (roots), Ipomoea batatas, Pangola 
grass roots, Passiflora foetida, rice stool, Leersia hexandra. This widespread 
and abundant root aphid often causes reddish-purple leaf discoloration. 
Heie (1967)—Kwalalessi, New Britain. Eastop (1966). 

57. Geoica lucifuga (Gehntner, 1897). 

BM; T—Goroka; Wau. 


Host PLAnt INDEX 

Albizia chinensis (Osb.) (Leguminosae-Mimosoideae)—A phis (Cerosipha) gossypii 
Glover. 

Amaranthus sp. (Amaranthaceae)—Toxoptera aurantu (B.d.F.). 

Arachis hypogaea li. (Leguminosae-Lotoideae)—Aphis (Pergandeida) craccivora 
Koch. 

Araucaria cunninghami Ait. (Araucariaceae)—Neophyllaphis araucariae Tak. 

Avocatlo—see Persea americana. 

Bamboo—see Bambusa sp. 

Bambusa sp. (Gramineae)—Pseudoregma bambusicola (Tak.). 

Banana—see Musa sp. 


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190 APHIDS OF PAPUA AND NEW GUINEA 


Bean—see Phaseolus, Vigna. 

Breynia cernua Muell. Arg. (Huphorbiaceae)—Schoutedenia lutea (v. d. G.). 

Cacao—see Theobroma. 

Caladium sp. (Araceae)—Pentalonia nigronervosa Coq. 

Camellia sinensis (L.) Kuntze (Theaceae)—Toxoptera aurantii (B.d.F.). 

Carica papaya L. (Caricaceae)—Aphis (Cerosipha) gossypii Glover. 

Cassia sp. (Leguminosae-Caesalpinioideae)—Aphis (Cerosipha) gossypii Glover. 

Casuarina equisetifolia Forst. (Casuarinaceae)—Lachnus tropicalis (v.d.G.). 

Celosia argentea (L.) Lautb. and K. Schum. (Amaranthaceae)—Hyalopterus pruni 
(Geoff.). 

Centrosema pubescens Benth. (Leguminosae-Lotoideae)—Aphis (Pergandeida) 

cracewora Koch. 

Chrysanthemum sp. (Compositae)—Macrosiphoniella (Pyrethromyzus) sanborni 
(Gillette). 

Citrus sp. (Rutaceae)—Toxoptera citricidus (Kirkaldy). 

Cocoa—see Theobroma cacao. 

Cocos nucifera Li. (Palmae)—Astegopteryx rappardi H.R.L., Cerataphis variabilis 
H.R.L. 

Coconut—see Cocos nucifera. 

Coffea arabica L. (Rubiaceae)—Toxoptera aurantii (B.d.F.). 

Coffee—see Coffea arabica. 

Colocasia sp. (Araceae)—Aphis (Cerosipha) gossypi Glover, Pentalonia nigro- 
nervosa Coq. 

Cowpea—see Vigna sinensis sinensis. 

Crotalaria anagyroides H.B. and K. (Leguminosae-Lotoideae)—Brachycaudus 
helichrysi (Kitb.), Myzus ornatus Laing, Myzus (Nectarosiphon) persicae 
(Sulz.). 

Cucurbita moschata Duchesne (Cucurbitaceae)—Aphis (Cerosipha) gossypii 
Glover. 

Cyphomandra sp. (Solanaceae)—Aphis (Cerosipha) gossypu Glover, Myzus 
(Nectarosiphon) persicae (Sulz.). 

Ficus sp. (Moraceae)—Greenidia ficicola Tak., 1921. 

Fig—see Ficus. 

Gladiolus sp. (Iridaceae)—Myzus (Nectarosiphon) persicae (Sulz.). 

Gliricidia sepium (Jacq.) Steud. (Leguminosae-Lotoideae)—A phis (Pergandeida) 
cracciwora Koch. 

Glycine max (L.) Merr. (Leguminosae-Lotoideae)—Aphis (Cerosipha) glycines 
Mats. 

Gramineae—Brachysiphoniella montana (v.d.G.), Ceratovacuna lanigera Zehnt., 
Longiunguis sacchari (Zehnt.), Macrosiphum (Sitobion) leelamaniae Kan. 
David, Pseudoregma bambusicola (Tak.), Rhopalosiphum maidis (Fitch), 
Tetraneura radicicola Strand, Tetraneura nigriabdominalis (Sas.). 

Grasses—see Gramineae. 

Hibiscus abelmoschus L. (Malvaceae)—Aphis (Cerosipha) gossypui Glov. 

Hibiscus rosa-sinensis L. (Malvaceae)—A phis (Cerosipha) gossypii Glover. 

Imperata cylindrica (L.) Beauv. (Gramineae)—Tetraneura radicicola Strand. 


Ipomoea batatas (L.) Lam. (Convolvulaceae)—Myzus (Nectarosiphon) persicae 
(Sulz.), Tetraneura nigriabdominalis (Sas.). 


Kunai grass—see Imperata cylindrica. 


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kK. P. LAMB 191 


Leersia hecandra Sw. (Gramineae)—Brachysiphoniella montana (v.d.G.), Tetra- 
neura nigriabdominalis (Sas.). 

Maize—see Zea mays. 

Malva parviflora L. (Malvaceae)—Aphis (Cerosipha) gossypu Glover. 

Musa spp. (Musaceae)—Pentalonia nigronervosa Coq. 

Oryza sativa Li. (Gramineae)—Tetraneura nigriabdominalis (Sas.). 


Passiflora foetida Li. (Passifloraceae)—Myzus (Nectarosiphon) persicae (Sulz.), 
Tetraneura nigriabdominalis (Sas.). 


Paw-paw—see Carica papaya. 

Peanut—see Arachis hypogaea. 

Pepper—see Piper sp. 

Pepper vine—see Vitis arborea. 

*Persea americana Mill. (Lauraceae)—Sinomegoura citricola (v.d.G.). 
Phaseolus sp. (Leguminosae-Lotoideae)—A phis (Pergandeida) craccivora Koch. 
Physalis peruviana L. (Solanaceae)—Neomyzus circumflecus (Buckt.). 

Piper sp. (Piperaceae)—Toxoptera aurantu (B.d.F.). 

‘* Pit-pit ’’—see Saccharum robustum. 

Platycertum wandae Raciborski (Polypodiaceae)—Micromyzus ? katoi (Tak.). 
Potato—see Solanum tuberosum. 

Potato, sweet—see Ipomoea batatas. 

Pumpkin, native—see Cucurbita moschata. 

Rice—see Oryza sativa. 

Rosa sp. (Rosaceae)—Macrosiphum rosae (L.). 


Saccharum robustum Brandes and Jesweit ex Grassl. (Gramineae)—Ceratovacuna 
lanigera Zebut. 


Saccharum officnarum L. (Gramineae)—Longiunguis sacchari (Zehnt.). 

Smilax sp. (Liliaceae)—Neomyzus circumflerus (Buckt.). 

Solanum tuberosum L. (Solanaceae)—Aphis (Cerosipha) gossypu Glover. 

Sorghum bicolor (lu.) Moench (Gramineae)—Rhopalosiphum maidis (Fitch). 

Soya bean—see Glycine max. 

Staghorn fern—see Platycerium. 

Sugar-cane—see Saccharum officinarum. 

Sweet potato—see Ipomoea batatas. 

Taro—see Colocasia sp. 

Tea—see Camellia sinensis. 

Tephrosia sp. (Leguminosae-Lotoideae)—Aphis (Cerosipha) gossypii Glover. 

Theobroma cacao lL. (Sterculiaceae)—Toxoptera auranti (B.d.F.), Cervaphis 
rappardi H.R.L., Cerataphis variabilis H.R.L. 

Tree tomato—see Cyphomandra. 

Vigna sinensis sesquipedalis (L.) Fruw. (Leguminosae-Lotoideae)—Aphis (Per- 
gandeida) craccivora Koch. 


Vigna sinensis sinensis Stickm. (Leguminosae-Lotoideae)—Aphis (Pergandeida) 
craccivora Koch. 


Vitis arborea (Vitaceae)—Myzus (Nectarosiphon) persicae (Sulz.), Toxoptera 
aurantit (B.de.F.). 


Yard-long bean—see Vigna sinensis sesquipedalis. 
Zea mays L. (Gramineae)—Rhopalosiphum maidis (Fitch). 


PROCEEDINGS OF THE LINNEAN Soctety oF New SoursH Watss, VoL. 96, Part 3 


192 APHIDS OF PAPUA AND NEW GUINEA 


ACKNOWLEDGEMENTS 
The author gratefully acknowledges the assistance of Dr. V. F. Hastop, 
British Museum (Natural History), and Dr. Mary Carver, Waite Agricultural 
Research Institute, for the identification of much material and for helpful 
discussions. Assistance in trapping was received from Miss K. Bevan and 
Messrs. G. O. Bedford, B. Gray and R. Hynes. I wish to thank Dr. P. Freeman, 
Keeper of Entomology, British Museum (Natural History), for giving me access 

to the aphid collections of the British Museum. 


References 


CaLLILuneG, V. J., 1967.—A faunistic study of Philippine aphids (Aphididae, Homoptera). The 
Philippine Agriculturist, 51: 93-170. 


Dun, G. S., 1954.—Annual report of the Senior Entomologist, Department of Agriculture, Stock 
and Fisheries, 1952-53. Papua N. Guin. Agr. Gaz., 8 (3): 18-27. 


Eastor, V. F., 1966.—A taxonomic study of Australian Aphidoidea (Homoptera). Aust. J. 
Zoology, 14: 399-592. 


Hein, O. E., 1967.—Aphids from the Philippines and the Bismarck Islands, with description 
of a new species of Greenideoida. Hnt. Medd., 35: 333-340. 


Hitte Ris LampBers, D., 1967.—On the genus Neophyllaphis Takahashi, 1920 (Homoptera, 
Aphididae) with descriptions of two new species. Zool. Meded., 42: 55-66. 


Szpent-Ivany, J. J. H., 1956.—New insect and host plant records in the Territory of Papua and 
New Guinea. P.N.G. Agric. J., 11 (3): 82-87. 


, 1958.—Insects of cultivated plants in the Central Highlands of New Guinea. Proc. X. 
Intern. Congr. Hnt., 3: 427-438. 


, 1961.—Insect pests of Theobroma cacao in the Territory of Papua and New Guinea. 
P.N.G. Agr. J., 13 (4): 127-147. 


Szpent-Ivany, J. J. H., and Cattry, A., 1960.—Host plant and distribution records of some 
insects in New Guinea and adjacent islands. Pacific Ins., 2 (3): 255-261. 


PROCEEDINGS OF THE LINNEAN SocreTy or New Souru Wates, Vou. 96, Part 3 


PHENOLOGY OF SPECIES COMMON TO THREE SEMI-ARID 
GRASSLANDS 


O. B. WILLIAMS* 


OSI.R.O., Division of Animal Physiology, The Ian Clunies Ross 
Animal Research Laboratory, Parramatta, N.S.W., 2150 


[Accepted for publication 19th May 1971] 


Synopsis 
Australian phenological studies are reviewed briefly before presenting data on the leafing 
phenology of seven species and the flowering phenology of 25 species common to (i) a Danthonia 
caespitosa grassland on Riverina clay, (11) a Stipa variabilis-D. caespitosa grassland on Billabong 
clay, and (i) a Stipa aristiglumis-D. caespitosa grassland on Deniboota loam. Data were 
collected for 1952 (annual rainfall 18-4 in.) and 1953 (annual rainfall 12-9in.). The disclimax 
grasslands adjoin on a level depositional plain situated in the south-west of New South Wales. 


The differences in leafing phenology of a species common to the three grasslands appear to 
be related to soil differences such as permanent wilting percentage (15-bar value) and water- 
holding capacity. This is more evident for warm-season species than for cool-season species. 


The growth-rhythms of the short-lived annual plants, which include many exotics, and the 
autumn- to spring-growing perennials indicate that the three disclimax grasslands are in 
equilibrium with the present annual climatic cycle. The remaining few perennial plants which 
respond to summer rainfall are related to, or are the indigenes of, the original communities from 
which the grasslands have developed. 


In those species which bloom once in the spring-early summer period, the onset of flowering 
appears to be related in part to day-length change. For most species the difference in the date 
of the onset of flowering between years in the one grassland appears to be greater than the 
differences between grasslands. Those species which have more than one flowering period do not 
exhibit a fixed relationship between day-length in spring-early summer, and day-length at the 
commencement of any other bloom period. 


It is concluded that the difference between the grasslands is found more in differences in the 
number of individual plants of each species and in the leafing phenology of the warm-season 
species common to the three grasslands, than in differences in species composition and in the 
flowering phenology of those species with only one bloom period. 


INTRODUCTION 


Moore (1880), in his Anniversary Address to the Royal Society of New South 
Wales, asked pastoralists to collect information on the habitat, vegetative and 
flowering performance of plant species on their sheep and cattle runs. No useful 
data appear to have been collected. Haviland (1886) compiled eight lists of 
indigenous species found about Sydney, N.S.W., and recorded the months in 
which they were in bloom. Maiden (1909) published a short bibliography of 
Australian phenological literature. 


Heyward (1931) constructed a phenological record for all 554 indigenous 
Victorian genera using herbarium records supplemented by personal observa- 
tions. She noted that most species did not vary their flowering dates by more 
than a few weeks even though temperature, rainfall, or elevation varied 
substantially. After discussing the importance of environmental factors in 
natural selection, and in particular the influence of the dry, hot summer, Heyward 
concluded that no single factor could be the sole determinant of time and duration 
of flowering. In this study the flowering periods of all species in a genus were 
combined to give a generalized picture. 


* Formerly C.S.1.R.O., Division of Plant Industry, Riverina Laboratory, Deniliquin, N.S.W., 
2710. 


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194 PHENOLOGY OF SPECIES COMMON TO SEMI-ARID GRASSLANDS 


Patton (1933) recorded the number of species flowering in each month on a 
heath at Cheltenham, Vic. He noted the large spring peak of bloom, that 
several indigenes performed exceptionally by flowering in summer, and that 
summer rainfall seldom elicited a plant response because of high evaporation. 
Patton (1936) recorded a similar late spring peak of bloom in an indigenous 
tussock grassland near Melbourne, Vic. 

Biddiscombe et al. (1954) reported leafing and flowering phenology of the 
major perennial grasses and winter-growing herbs in a disclimax grassland at 
Trangie, N.S.W. The first detailed list of flowering dates for individual species 
in an Australian community was reported by Specht and Rayson (1957) for the 
heath near Keith, S.A. Derrick (1962) presented charts for all Australian States 
(excluding the Northern Territory) which show the approximate flowering 
season of Many common indigenes and exotics. Groves and Specht (1965) 
have presented flowering times for two areas of heath vegetation at Wilson’s 
Promontory, Vic., and Groves (1965b) for indigenous and naturalized species 
in a Themeda tussock grassland at St. Albans, Vic. 

The first report on both flowering and leafing phenology for more than one 
year was presented by Williams (1961) for a number of species in a Danthonia 
caespitosa grassland on Riverina clay at Deniliquin, N.S.W. In the present 
paper the 1952 and 1953 data on leafing and flowering phenology of selected 
species in this grassland are compared with the same species in two adjoining 
semi-arid grasslands, Stipa variabilis-D. caespitosa on Billabong clay, and 
S. aristiglumis-D. caespitosa on Deniboota loam. ‘The data are considered in 
relation to estimates of the periods during which soil water is available for plant 
growth. Differences and similarities in phenology are discussed. Climate is 
common to all three communities because they adjoin, trees are absent, short- 
grasses and herbs predominate, and the general topography is featureless. 


ENVIRONMENT 
Climate 

Detailed characteristics of the Deniliquin climate have been presented 
previously (Williams, 1956). The seasons are taken as summer (December to 
February), autumn (March to May), winter (June to August), and spring 
(September to November). The mean annual rainfall is 15-8in., and cool- 
season rains are more reliable than summer rains. 

Rainfall recorded in 1952 and 1953 at the meteorological station proximate 
to the three sites is set out in the lower part of Fig. 1 as in./15 days. In 
1952 the annual rainfall was 18-4 in., or 10°% above average, while in 1953 it 
was 12-9in., 23°% below average. 

Summers are hot, with occasional thunderstorms. January is the hottest 
month, with a mean maximum temperature of 89-5° F. and a mean minimum 
of 61° F. From January to March, 1952, there were 34 days, and from October 
to December eight days with maximum temperatures in excess of 90° F. The 
corresponding values in 1953 were 33 and 14 days. 

Winters are cool. In both 1952 and 1953 there were approximately 30 
light frosts (screen temperature >32° <36° F.) and 16 severe frosts (screen 
temperature <32° F.). 

Day length (civil twilight) at the experimental site ranges from a maximum 
of 15 hours in December to a minimum of 10 hours in June. 

In summary, incidence and quantity of rainfall and the period of heavy 
frost were the factors which differed most markedly between the two years. 


Soils 
(i) Topography and general distribution 

The topography and soils of the general area have been described by 
Churchward (1958). The three experimental sites are located at the C.8.I.R.O. 


PROCEEDINGS OF THE LINNEAN SoOcIETY OF NEw SoutH WALES, VoL. 96, Part 3 


O. B. WILLIAMS 195 


Falkiner Memorial Field Station, the D. caespitosa and Stipa variabilis-D. 
caespitosa grasslands on the lower and upper levels of the clay plains, and the 
S. aristiglumis-D. caespitosa grassland on the loam of the Stud Park River 
Ridge. 


(ii) Soil-types 

The soils at each site are similar to those described and named by Smith 
(1945), Butler and Johnston (1946), and Johnston (1953) as Riverina clay, 
Billabong clay, and Deniboota loam. 


Riverina clay is a grey clay (Stace et al., 1968, p. 79) with a 0-0-5in. A 
horizon of slightly structured light clay which is hard when dry and sticky when 
wet. The B, horizon of 0-5—10 in. is a moderately structured heavy clay over- 
lying grey poorly-structured heavy clay. The permanent to semi-permanent: 
cleavage planes extend to the surface. All three components of the gilgai 
micro-relief, the puff, shelf and depression are represented (Williams, 1955). 


Billabong clay is a brown clay (Stace et al., 1968, pp. 79, 103) with a 0-0-5 in. 
A horizon of grey-brown compacted clay loam which is hard when dry and sticky 
when wet. The B, horizon of 0-5—-12 in. is a dark grey-brown massive heavy 
clay with blocky structure. It is hard when dry and intractable when wet. 
Below 12 in. the soil is a poorly-structured heavy clay. Cleavage planes are 
few and rarely extend to the surface. There are few areas of puff, and a small 
amount of depression. 


Deniboota loam is a red-brown earth (Stace et al., 1968, pp. 211, 214) with 
a 0-3 in. A, horizon of brown loam which is hard when dry and friable when. 
wet. The A, horizon is 3-6 in. of grey-brown sandy clay loam which is slightly 
compact and hard when dry but friable when wet. The B, and B, horizons 
from 6—24 in. are medium clays with well developed medium prismatic structure 
grading into grey-brown clay. : 

The main chemical differences between the three soils in the top 24 in. are 
(1) the acid reaction of the Billabong clay and Deniboota loam surfaces compared 
with the alkaline reaction of the Riverina clay (pH 6-1, 6-5 v. 7-5) and the 
pronounced alkaline reaction (pH 8-8) of the loam below 9 in., (2) the small 
amount of chloride in the Deniboota loam profile compared with the amounts 
in the Riverina and Billabong clays (0-034 v. 0-237 and 0-206 at the 9-18 in. 
depth, expressed as NaCl per cent. oven dry weight). 


The differences in nitrogen and phosphorus status between the various. 
surface soils are small, with a range of 0-052 to 0-100% and 0-017 to 0-020°% 
respectively. 


Both calcium and magnesium on the exchange complex of Riverina clay 
greatly exceed the values obtained from Deniboota loam (13-98 v. 2-27 and 
14-05 v. 1-92 m-equiv./100 g. oven dry weight). 


(iii) Soil moisture relationships 


The periods over which water in the 0-3 and 3-9 in. depth is available for 
plant growth are set out in the lower part of Fig. 1, together with a record of 
the sampling dates and the incidence of rain sufficient to raise the moisture 
content of the 0-3 in. depth to the 15-bar level and above for short periods 
between sampling dates. These periods were calculated for the various depths. 
by gravimetric determinations of soil moisture for two sites in each gilgai 
component on the clay soils, and at two sites in the loam, 15-bar values for each 
depth, and bulk densities for the soil horizons. Slivers of wet soil were taken 
from the sides of cracks after rain and the soil water content determined. 


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196 PHENOLOGY OF SPECIES COMMON TO SEMI-ARID GRASSLANDS 


The amount of water which raised the moisture content of the surface 0-3 in. 
from the minimum field value to the 15-bar value ranged from 0.08 in. for 
Deniboota loam, 0-28 and 0-40 in. for the shelf of Riverina clay and Billabong 
clay, to 0-50 and 0-56 in. for the depression of the Riverina clay and Billabong 
clay. Small amounts of rain are sufficient to provide available water in the 
surface of Deniboota loam. 

The available water within the maximum wetted profile over the 1952-53 
period ranged from 4-3in. in the Deniboota loam, 1-7 and 1-0in. in the 
depression and shelf of the Riverina clay, to 0-8 and 0-4 in. in the depression 
and shelf of Billabong clay. 

In 1952 the Deniboota loam had available water in the 0-3 in. depth for 
270 days, the Riverina clay for 260 days (depression) and 240 days (shelf), the 
Billabong clay for 220 days (depression) and 140 days (shelf). In 1953 these 
periods were shortened by from 70 to 110 days. 

Depressions tend to be wetter than 15-bar values, per se, indicate because 
depressions are recipients of run-off from the massive surface of the neighbouring 
shelf. Further, run-off into cleavage planes modifies the generalized available 
water situation because it produces localized pockets of wet soil at depth. Water 
penetration is more uniform in Deniboota loam. 


Vegetation 
The three grasslands appear to differ markedly from the original communities 
of 100 years ago. 


(i) Danthonia caespitosa grassland. 

This grassland has been the subject of several papers (Williams, 1955, 1956, 
1961). It has developed from an Atriplex nummularia Lindl. shrub-steppe 
community, possibly by way of an A. vesicaria Hew. ex Benth. community. 

The last A. vesicaria plants were eliminated from the experimental area by 
sheep grazing in 1948. 


(ii) Stipa variabilis-Danthonia caespitosa grassland. 

This grassland appears to have developed from the Acacia pendula- Atriplex 
nummularia association (Moore, 1953a, pp. 514-5). Few areas of unmodified 
shrub woodland now exist. 

The present grassland has been described by Moore (1953), p. 561) as a 
disclimax community, and Williams (1956) has measured its botanical com- 
position. Of the original dominants, only scattered plants of Kochia aphylla 
R.Br. remain. 


(iii) Stipa aristiglumis-Danthonia caespitosa grassland 


This grassland is a disclimax community at the western margin of the 
Eucalyptus woollsiana-Callitris columellaris association within the H. woollsiana 
alliance (Moore, 1953a, p. 508). Small areas of the original savanna woodland 
still exist to the east of the Field Station. 


The present grassland developed under intensive sheep grazing and the 

felling of trees for drought-fodder, structural timber or firewood. 
PROCEDURE ; 

The procedure followed the pattern of the previous study (Williams, 1961), 
with the frequent recording of species behaviour within a fixed area in each of 
the three grasslands. These observations were made monthly, or more 
frequently, e.g. every two or three days, when plants were flowering. 


The commencement of leafing, onset and duration of flowering and leaf 
senescence were recorded for each species. The periods when plants were 


PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WaLEs, Vow. 96, Part 3 


O. B. WILLIAMS 197 


vigorously vegetative, or were making limited leaf growth were estimated using 
plant height, plant diameter, or leaf length, appropriate to the growth habit of 
the particular species. Recording commenced in mid-1951 and continued 
until the end of 1954. Confirmatory checks of bloom periods were made between 
1956 and 1961. 

Soil sampling was started in mid-1951 and continued until the end of 1953. 
The two years 1952 and 1953 with coincidental vegetation and soil moisture 
data are used in this paper. 


RESULTS 
The phenological data are presented in two sections: (a) the leafing 
phenology of seven species common to all three grasslands in 1952 and 1953, 
and (b) the flowering phenology for 25 common species during the same period. 


(a) Leafing Phenology 

Figure 1 illustrates the leafing phenology of seven species chosen because 
there were complete records for both soils and years and because they illustrate 
the behaviour of many species in the grasslands. Rainfall, frost periods and 
periods of available water in the 0-3 and 3-9 in. depths for each site are shown 
for 1952 and 1953. 


ARGUE SPECIES GRASSLAND 1952 1953 


1.—A HORDEUM LEPORINUM LINK 


{" STIPA VARIABILIS HUGHES 
2 


P DANTHONIA CAESPITOSA GAUDICH. (+ 


P ATRIPLEX SEMIBACCATA R BR 


P SIDA CORRUGATA LINDL 


P CHLORIS ACICULARIS LINDL 


A-B CHLORIS TRUNCATA R BR 


| 36° OR CESS 


—_ 
FROST PERIOD 32° OR LESS ae OR LESS 


}___—___->"__, = 
DEPRESSION >| a ooo —EE ° GE 
SHELF . om mE «COO (O00 mo Ommo oc 0 6(0CamIE EEE 
Go tT cle la SHELF or CO 0 0 mets 0 9 COOH © ° miteatbetlios © ES a 
° 


SOIL SAMPLING DATES 


(IN. 7 15 DAYS) 


4 
3 
RAINFALL = aL. 
, 
i) 


Fig. 1. 


The three groups are numbered 1, 2, and 4, and are arranged as previously 
described (Williams, 1961) in the order of predominantly cool-season growth, 
through growth in all seasons, to predominantly warm-season growth. Groups 
3 and 5 are omitted because they contained no species which grew continuously 
on all three soils in each year. 


The difference between the sites is illustrated by the performance of species 
as follows : Deniboota loam and the depressions in Riverina clay tend to maintain 
green leaf longer, as with Hordeum leporinum in spring and early summer, than 
do the other soils. In Group 2-1952 Stipa variabilis tends to start growth first 
in Deniboota loam, then Billabong clay, and Riverina clay. In Group 4 there 
were differences in species behaviour on the three soils, e.g. Atriplex semibaccata 
(May, 1953), Sida corrugata (January-February, 1952), Chloris acicularis (March, 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WaAtEs, Vout. 96, Part 3 


198 PHENOLOGY OF SPECIES COMMON TO SEMI-ARID GRASSLANDS 


D. CAESPITOSA S. VARIABILIS D. CAESPITOSA 
D. CAESPITOSA S. ARISTIGLUMIS 


No OF SPECIES FLOWERING 
—) nN 
ts} iS 

q 

i 

- 

i= 

- 


ft si) taf Est 4 1 1 4 ut | 
TLiELIMCRAGIN, SGPURUIGEA EGF UOMINID Blas SU FIEINIt AUR SRA MIS? 70) SR UES Mi TALIM) Ue) SAW iSiaOeN SD: 
Fig. 2. 
SPECIES GRASSLAND 
YEAR |\yilude JESU WA Maa yA 8S) SOM NID 
1952. ge 
52 ° | = 
As ERODIUM CICUTARIUM (L.) L'HERIT. ecg L 
53 e 
ee0o _— 
A® VULPIA BROMOIDES (L.)S.F. GRAY oe oes i aa | 
e = = 
A HELIPTERUM PYGMAEUM (BENTH) DC 2 are =a ca 
Doane = 
A HELIPTERUM MOLLE (A. CUNN. ex DC ) S28 | | | eee] | 
PG. WILSON 33 Re E-= 
es 
A® BROMUS RUBENS L. 52 eSs 
53 683 
A  HELIPTERUM AUSTRALE (A. GRAY) DRUCE 22 ine =e 
ie 
A® HORDEUM LEPORINUM LINK. ae oss See 
53 
2 
A  ASPERULA CONFERTA HOOK. f 22 ES 
53 ae 
P  KOCHIA EXCAVATA JM BLACK oa 
53 : 
e 
A ERODIUM CRINITUM CAROLIN oe 
53 
52 
A  DAUCUS GLOCHIDIATUS (LABILL ) 538 eee 
FISCH, MEY. & AVE-LALL of? 
Az MEDICAGO POLYMORPHA L Ss iadlnal al PI 
Eat = 
52 ee ; 
P  STIPA VARIABILIS HUGHES ae 
53 e : 
ec0e 
A® HEDYPNOIS CRETICA (L.) WILLD pe aaae 
Bute 
OOK 
P  PTILOTUS SPATHULATUS (R. BR.) POIR ena re mu 
53 os$ rel 
A® LOLIUM RIGIDUM GAUDICH 92 “33 
53 
o$3 
52 ° 
P DANTHONIA CAESPITOSA GAUDICH ots 
53 ee 7 
eee 
Joe LIM VyAl IMs (9D JO TAe SH Or eN@ fb 
Fig. 3. 


PROCEEDINGS OF THE LINNEAN Society or New SourH WALES, VOL. 96, Part 3 


O. B. WILLIAMS 199 


1952), but more particularly C. truncata (January to April, 1952; August to 
October, 1952; and September to November, 1953). 

The growth differences between 1952 and 1953 were substantial. In 1952 
active growth commenced in mid-March, but in 1953 there were two false starts, 
and the species in Group 1 did not commence active growth until the end of June. 
The heavy rainfall in October, 1952, with the vigorous leaf-growth of Group 4 
species, which continued until the end of December, contrasts strongly with the 
tapering off in the growth of these species at the conclusion of the 1953 spring. 
Low winter temperatures and frosts do not prohibit leaf growth in Groups 1 
and 2. 


Years in which the soil moisture is low in autumn and winter, as in 1953, 
are not unusual. 


(b) Flowering Phenology 
(i) Flowering rhythms 
In Fig. 2 the flowering rhythms of the three grasslands are set out. The 


maximum number of species observed flowering in each particular month of the 
two-years experimental period has been used to construct this figure. 


A feature of these flowering rhythms is the contribution made by species 
that have more than one bloom period. 


SPECIES GRASSLAND 
YEAR 
1952 48 
53 
52 
53 
52 
53 
52 
53 
52 
53 
52 
53 
52 
53 


P HELICHRYSUM APICULATUM (LABILL, DC. 


A WAHLENBERGIA GRACILENTA LOTHIAN 


A-B CHLORIS TRUNCATA R. BR 


P GOODENIA SUBINTEGRA F. MUELL. 


P CHLORIS ACICULARIS LINDL. 


P ATRIPLEX SEMIBACCATA R. BR. 


P SIDA CORRUGATA LINDL. 


(ii) Flowering dates 


The flowering dates for species common to the three grasslands are set out 
in Figs 3 and 4. 


Figure 3 summarizes the records for 18 species which have consistently 
exhibited one main bloom period between July and December, but at no other 
time. They are arranged in order of flowering. Most plants in these disclimax 
grasslands belong to this category. 


The onset of the main bloom period differed between soils for Ptilotus 
spathulatus in 1952, Helipterum pygmaeum in 1953, and to a lesser degree for 
Helupterum molle and Daucus glochidiatus in 1953. It differed between years 
for Helipterum australe and Hordeum leporinum. Though the bloom periods of 
some species, e.g. Hordeum leporinum started earlier in 1952 than in 1953, other 


PROCEEDINGS OF THE LINNEAN SocrEetTy oF New SoutH Watss, Vou. 96, Part 3 


200 PHENOLOGY OF SPECIES COMMON TO SEMI-ARID GRASSLANDS 


species, e.g. Vulpia bromoides, behaved in the opposite manner. Peak bloom in 
1952 was generally a few days to a week earlier than in 1953. 


Species which may exhibit one bloom period between July and December, 
and another in summer and autumn, cause the February to April peak in Figure 2. 
The flowering periods of seven species in this group, arranged in order of spring 
flower appearance, are shown in Fig. 4. 


The onset of the spring bloom period differed between years for Wahlenbergia 
gracilenta and Chloris truncata. Differences between the soils in either 1952 or 
1953 were seldom greater than one week for these species, but up to six weeks 
for species such as Goodenia subintegra. 


Summer and autumn flowering varies markedly between years, and, with 
the exception of Chloris acicularis, between soils in the one year. 


DISCUSSION 
Leafing Phenology 


Differences in leafing phenology of common species in the three grasslands 
are apparent in early 1952 in the S. aristiglumis-D. caespitosa grassland, where 
there was a tendency for Group 1 species to start growth first and for the Group 4 
species to Maintain vigorous growth over a longer period than in the other two 
grasslands. This performance was repeated in late 1952, but not in late 1953. 


The summer-growing species of Group 4 are rather irregular in their 
performance, because they depend on the erratic summer rains. Even so, 
there are several important differences between the grasslands for species such as 
Sida corrugata and Chloris truncata. 


These differences are attributed to differences in soil water between 
Deniboota loam, Billabong clay and Riverina clay, and in particular, to the low 
15-bar value, high permeability, and large water-holding capacity of Deniboota 
loam relative to the other soils. The crude nature of the relationship between 
the overall leafing phenology and available water in Fig. 1 reflects the 
inadequacy of the gravimetric method for determining moisture in soils which 
rains penetrate unevenly and roots do not exploit uniformly. The relationship 
between the drying cycles and the leafing phenology is particularly unsatis- 
factory. Itis noteworthy that the more successful studies of this type have been 
recorded on permeable soils in regions of pronounced seasonal rainfall (Specht, 
1957): 


The pattern of vegetative growth in 1952 and 1953 illustrates the performance 
in a long and a short season ; performances which occur frequently over large 
tracts of southern Australia. There is nothing atypical in the two false starts 
of the 1953 season and the associated contrast in leafing phenology between 
annuals and perennials. Heavy rain makes moisture available for germination 
or growth on all three soils, whereas a light rain elicits a response only from the 
loam soil. 


Specht and Rayson (1957), in their studies of the vegetative performance 
of heath species at Dark Island, South Australia, reported that, as expected 
in southern Australia, the geophytes made maximum growth in spring. However, 
some perennial components of the heath grew during hot, dry summers on soil 
water stored from the preceding winter-spring rains. Occasional ineffective 
rain, or rare effective storms between December and March, do not appear to 
upset the notion that this lack of a relationship between the vegetative growth 
cycle of the dominant perennial (Banksia ornata), and the present rainfall regime 
is due to a long-term shift in climate (Specht and Rayson, 1957 ; Specht, 1958). 
These summer-growing species Maintain vegetative growth patterns which are 
possibly characteristic of this previous climate. Burbidge (1960) has commented 


PROCEEDINGS OF THE LINNEAN Society or NEw SourH WALES, Vou. 96, Part 3 


O. B. WILLIAMS 201 


upon this out-of-phase characteristic for Eucalyptus and Banksia spp. in Western 
Australia, and Johnston and Briggs (1963) for Proteaceae. It is likely that these 
rare effective storms, together with impoverished soils, frequent burning, and the 
low intensity of grazing by introduced herbivores have been important in main- 
taining the dominants at Dark Island. 

Groves (1965a) has recorded the maximum growth-rate of Victorian heaths 
in summer, from December to March, and of Themeda tussock grassland (Groves, 
1965b) from October to early December. He contrasted these performances 
with the performance of a sown pasture of Lolium perenne and Trifolium sub- 
terraneum, which exhibited a maximum growth-rate from August to September 
(Twentyman, 1938). Groves commented further that the productivity of the 
Danthonia spp. in the Themeda grassland suggests that a Danthonia grassland 
may reach its maximum growth-rate some months before the Themeda com- 
munity. Within most communities studied so far there are species groups 
which do not conform to the general community pattern. For example, Holland 
(1968), in seasonal growth studies in mallee communities in central New South 
Wales and western Victoria, has shown that tall shrubs and mallee eucalypts 
do make summer growth, but field layer plants do not. The three grasslands 
in the present study contain a wide range of phenological performances which 
similarly preclude a statement as to the particular community growth-rhythm. 

In the Riverine plain the original communities appear to have been 
dominated by species with potential growth-rhythms of the warm-season type. 
From 1845 onwards these communities have been shaped by the process of 
species elimination, species invasion under the influence of sheep, rabbits, various 
other herbivores, and intermittent droughts (Williams, 1968). These processes 
have favoured the plants that can complete their life-cycle within the more 
reliable rainfall period from May to October, resulting in either the dominance 
of indigenes which can cope with the new conditions or the dominance of annual 
and perennial species of the Mediterranean type, or a mixture of the two groups. 
The remaining species are wholly, or partly, opportunist and rely on the erratic 
summer rains. Amongst the wholly. opportunist species is Chloris truncata. 
Perennial species such as Chloris acicularis represent the vegetative pattern 
of the type of indigene which dominated the original communities. In spite 
of the wide-ranging root systems of these perennial indigenes, the growth pattern 
is not as regular as that of the heath dominants described by Specht and Rayson 
(loc. cit.), doubtless because soil moisture storage and rooting depth, even in 
Deniboota loam, are limiting on more occasions than they appear to be under 
heath vegetation. Thus each of the three communities expresses a wide range 
of species phenology. The exact form of this vegetative phenology is influenced 
by micro-topography, soil water differences between the grasslands, and different 
rainfall between years. 


Flowering Rhythms 

The flowering rhythms of the two species groups in each of the three 
grasslands are similar (Fig. 2). They appear to have counterparts in Themeda 
grassland (Groves, 1965b) and in southern heaths (Specht and Rayson, 1957 ; 
Groves and Specht, 1965), but not in montane communities in New Zealand 
(Scott, 1960; Clarke, 1968). 

In the Riverine plain this two-population situation probably reflects the 
partial species elimination and concurrent invasion of species discussed previously. 


Flowering Phenology 

It has already been suggested (Williams, 1961) that the regular onset of 
bloom for spring flowering species of the Danthonia caespitosa grassland may be 
attributed in part to changes in day-length. This suggestion appears to be 
valid for these species in the other two grasslands. 


PROCEEDINGS OF THE LINNEAN SocrEety or NEw SourH Watss, Vou. 96, Part 3 


202 PHENOLOGY OF SPECIES COMMON TO SEMI-ARID GRASSLANDS 


Plasticity in vegetative characteristics does not appear to extend to flowering 
phenology. At best, a large vegetative response may promote a small flowering 
response in bloom period, but no shift in date of first bloom has been noted so far. 
Extended, or additional bloom periods in such annual species as Medicago 
polymorpha and the two Hrodium spp. appear to be due to progressive flowering 
in favoured sites and the presence of more than one age group in the population. 

The processes of species elimination and invasion, coupled with the reliable 
rainfall period in winter and spring, have favoured annual or perennial species 
of the Mediterranean type. By seeding profusely before the end of October, 
with or without hard-seed production, these species maintain themselves in the 
community. 


The species which can flower in spring-summer and at other times are 
ephemerals of the Wahlenbergia gracilenta type or long-lived perennials of the 
Chloris acicularis type. An examination of the dates of first bloom has shown 
that the spring and autumn photoperiodic range for individual species is wide 
under natural conditions, and critical investigations using clones, transplants 
and field irrigation are needed. 


The frequent-flowering perennials at Deniliquin are important from the 
ecological viewpoint because they are either remnants of the indigenes which 
dominated the original communities or are similar to them in leafing and flowering 
phenology. Their population density is low, as might be anticipated in a pastoral 
zone. This also appears to be the case at Trangie (Biddiscombe, 1953). 


In principle, the maintenance of a plant in a community depends upon 
the longevity of the plants, the opportunity during that life-span to produce 
viable seed, the viability of the seed in the soil, and the opportunity to germinate 
and establish. Flowering, per se, may not mean much for trees and long-lived 
shrubs, but will mean a great deal for annuals (Holland, 1968). It will be 
critical in the early stage of colonization by a new species, becoming less important 
as the seed supply in the soil builds up and the area colonized increases. 


Until the advent of domestic stock, there seems to have been scant advantage 
for the indigenes to produce seed or to possess superlative seedling vigour, because 
the parent plants were long-lived. In contrast, today there is a survival 
advantage in heavy spring seeding, in leaf-growth during the reliable rainfall 
period, and possibly in a limited vegetative response to out-of-season rainfall. 
Although the grazing pressures applied to the communities should favour earlier 
flowering through changes in botanical composition, the erratic summer storms 
support a counter movement. Herein lies one more reason for the difficulty 
in manipulating grasslands of this type by grazing, per se (Williams, 1969). 

It is concluded that differences between species common to the three grass- 
lands are found to some degree in leafing phenology, particularly the leafing 
of the warm-season species, and to a more limited degree in the flowering 
phenology. The substantial difference between grasslands is not found so much 
in representation by different species or in the flowering phenology of those 
species common to the grasslands, as in the number of individuals of each species, 
and the leafing phenology of the warm-season species common to the three 
grasslands. Apparently the general soil differences, though substantial, have 
not been sufficient to provide a stimulus for ecotypic differentiation in the presence 
of the grazing animal over the past 100 years. 


References 


BippiscoMsE, EH. F., 1953.—A survey of the natural pastures of the Trangie District, New South 
Wales, with particular reference to the grazing factor. Aust. J. agric. Res.,4: 1-28. 


BrppiscoMsBE, EH. F., CurHsBertson, E. G., and Hurcnuines, R. J., 1954.—Autecology of some 
natural pasture species at Trangie, N.S.W. Aust. J. Bot., 2: 69-98. 


BursipGe, Nancy T., 1960.—The phytogeography of the Australian region. Awsi. J. Bot., 
8: 75-212. 


PROCEEDINGS OF THE LINNEAN Society oF NEw South Wates, Vou. 96, Part 3 


O. B. WILLIAMS 203 


Butuer, B. E., and Jounston, E. J., 1946.—A report on a detailed soil survey of Falkiner 
Memorial Field Station. Coun. Sci. Industr. Res. Aust. Div. Soils Rep. (Mimeo.) 

CHURCHWARD, H. M., 1958.—The soils and land use of the Denimein Irrigation District, New 
South Wales. C.S.JI.R.O. Aust. Soils and Land Use Ser., No. 27. 

CuaRKE, C. M. H., 1968.—Flowering periods of alpine plants at Cupola Basin, Nelson, New 
Zealand. N.Z. J. Bot., 6: 205-220. 

Derrick, Epiru A., 1962.—Relative importance of various plants in causation of hayfever and 
asthma in Australia. Med. J. Aust., 1: 972-977. 

Groves, R. H., 1965a.—Growth of heath vegetation. II. The seasonal growth of a heath on a 
ground-water podzol at Wilson’s Promontory, Victoria. Aust. J. Bot., 13: 281-289. 

, 1965b.—Growth of Themeda australis tussock grassland at St. Albans, Victoria. Aust. 
J. Bot., 13: 291-302. 

Groves, R. H., and Specut, R. L., 1965.—Growth of heath vegetation. I. Annual growth 
curves of two heath ecosystems in Australia. Aust. J. Bot., 13: 261-280. 

Havitannd, E., 1886.—Flowering seasons of Australian plants. Proc. Liyn. Soc. N.S.W., 11: 
1049-1051. 

HEYWARD, JEAN, 1931.—Flowering periods of Victorian plants. Proc. roy. Soc. Vic., 43 (n.s.) : 
154-165. 

Houanpd, P. G., 1968.—Seasonal growth of field layer plants in two stands of mallee vegetation. 
Aust. J. Bot., 16: 615-622. 

Jounston, E. J., 1953——Pedology of the Deniboota Irrigation District, New South Wales. 
C.S.I.R.O. Aust. Soils Publ., No. 1. 

JoHNSoN, L. A. S., and Brieas, BarBara G., 1963.—Evolution in the Proteaceae. Aust. J 
Bot. VW: 2I=61-. 

Marpen, J. H., 1909.—A plea for the study of phenological phenomena in Australia. J. Proc. 
roy. Soc. N.S.W., 43: 157-170. 

Moore, C., 1880.—Anniversary address. J. Proc. roy. Soc. N.S.W., 14: 1-18. 

Moors, C. W. E., 1953a.—The vegetation of the south-eastern Riverina, New South Wales. 
I. The climax communities. Avwst. J. Bot., 1: 485-547. 

, 1953b.—The vegetation of the south-eastern Riverina, New South Wales. II. The 
disclimax communities. Aust. J. Bot., 1: 548-567. 

Patton, R. T., 1933.—Ecological studies in Victoria—The Cheltenham flora. Proc. roy. Soc. 
Vic., 45 (n.s.): 205-218. 

, 1936.—Ecological studies in Vietoria. IV. Basalt plaims association. Proc. roy. 
Soc. Vic., 48 (n.s.): 172-191. 

Scorr, D., 1960.—Seasonal behaviour of some montane species. N.Z. J. Sci., 3: 694-699. 

Smith, R., 1945.—Soils of the Berriquin Irrigation District, N.S.W. Coun. Sci. Industr. Res. 
Aust. Bull., No. 189. 

Specat, R. L., 1957.—Dark Island Heath (Ninety-Mile Plain, South Australia). V. The water 
relationships in heath vegetation and pastures on the Makin sand. Aust. J. Bot., 5: 
151-172. 

, 1958. The geographical relationships of the flora of Arnhem Land. Amer.-Aust. 
Sci. Hupedit. to Arnhem Land, 3: 415-442. 

SpecHt, R. L., and Rayson, Parricia, 1957.—Dark Island Heath (Ninety-Mile Plain, South 
Australia). J. Definition of the ecosystem. Aust. J. Bot., 5: 52-85. 

Stacr, H. C. T., Hussiez, G. D., Brewer, R., et al., 1968.—A Handbook of Australian Soils. 
Rellim. Tech. Publ., Glenside, South Australia. 

TWENTYMAN, R. L., 1938.—Pasture improvement. J. Dept. Agric. Vic., 36: 155-170, 197-204. 

Wrttiams, O. B., 1955.—Studies in the ecology of the Riverine Plain. I. The gilgai microrelief 
and associated flora. Aust. J. Bot., 3: 99-112. 

, 1956.—Studies in the ecology of the Riverine Plain. II. Plant-soil relationships in 
three semi-arid grasslands. Aust. J. agric. Res., 7: 127-139. 

, 1961.—Studies in the ecology of the Riverine Plain. III. Phenology of a Danthonia 
caespitosa Gaudich. grassland. Aust. J. agric. Res., 12: 247-259. 

, 1968.—Studies in the ecology of the Riverine Plain. IV. Basal area and density 
changes of Danthonia caespitosa Gaudich. in a natural pasture grazed by sheep. Aust. J. 
Bot., 16: 565-578. 

, 1969.—Studies in the ecology of the Riverine Plain. V. Plant density response of 
species in a Danthonia caespitosa grassland to 16 years of grazing by Merino sheep. Aust. J. 
Bot., 17: 255-268. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wates, Vou. 96, Part 3 


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PROCEEDINGS OF THE LINNEAN SocrETy oF NEw SoutH Wars, Vot. 96, Part 3 


AUSTRALASIAN MEDICAL PUBLISHING CO. LTD. 
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CHECK LIST OF NORTHERN TERRITORY PLANTS 


G. M. CHIPPENDALE 
Forest Research Institute, Forestry and Timber Bureau, Canberra 


[Accepted for publication 23rd June 1971] 


Synopsis 
The need for a check list of Northern Territory plant species is discussed with a brief outline 
of the vegetation. New combinations are made for Borrerta involucrata (O. Schwarz) Chippendale 
and Canthium schultz (O. Schwarz) Chippendale. Tables show the geographic distribution of 
species, families, and of naturalized and adventive specics. The check list contains the names 
of 2,695 native species and 113 introduced species. A bibliography provides references to support 
the check list. 


INTRODUCTION 


The Northern Territory of Australia was little known when Ewart and 
Davies (1917) compiled ‘‘ The Flora of the Northern Territory ”’, a work which 
had many faults mentioned in a critical review (Anon., 1919). Not surprisingly, 
Specht (1958) for the northern area, and Chippendale (1959) for the central area 
listed many new records. As remarked by Cochrane (1967), the Northern 
Territory is “... still vegetatively relatively unexplored...”. Thus, a check 
list of the known species should be a useful basis for the present and future 
taxonomy and ecology in the area. 


This enumeration of the plant species occurring in the Northern Territory 
was compiled mainly while the author was working in the Herbarium of the 
Northern Territory at Alice Springs. However, in addition to using specimens 
in that herbarium, much relevant literature has been searched and is mentioned 
in the bibliography. Each species included is supported by a specimen seen by 
the author in an Australian herbarium or by the citation of a specimen in published 
work. Actual references to specimens are not given. 


Of the literature searched, some of the older references were inaccurate or 
vague. For instance, Bentham (1863-78) gave many records from Northern 
Territory locaJities and a few specifically from Central Australian localities, but 
his records from ‘‘ Northern Australia ’’ and ‘‘ Islands of the Gulf of Carpentaria ”’ 
were often wider, geographically, than the Northern Territory. Likewise, Ewart 
and Davies (1917) gave records from ‘‘ North Australia’’. Such records have 
not necessarily been accepted as occurring in the Northern Territory ; usually 
these species are included in the check list only if they have since been reliably 
collected. 


VEGETATION 


The four major pastoral districts of the Northern Territory (Fig. 1) are used 
to give an indication of the geographic occurrence of species. These districts 
are primarily administrative in origin, but broadly each represents different 
ecological conditions. 

The Darwin and Gulf District is dominated by tall open forest, predominantly 
of Hucalyptus tetrodonta and EF. miniata in the more northern parts, with an 
understorey of tall perennial grasses. Some shrubland has dense stands of the 
palm Livistona humilis which merges with communities including Calytriz, 
Grevillea, Persoonia, Acacia and Bossiaea. There are minor areas of monsoon 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wats, Vor. 96, Part 4 


208 NORTHERN TERRITORY PLANTS 


forest, and the seasonally flooded sub-coastal plains support grassland, sometimes 
surrounded by a narrow stand of tall Melaleuca leucadendron. Mangroves and 
beach dune species occur on the coastal strand. 


Open forests and woodland cover much of the Victoria River District, and 
include many species of Hucalyptus, with associated coarse perennial grasses. 
Erythrophloeum chlorostachys is prominent in some of these woodlands, and 
Acacia shirleyi forms a dense low forest in the eastern part of the district. Lesser 
areas are dominated by shrublands of Macropteranthes kekwickiit and Hacaecaria 
parvifolia. Grasslands composed mainly of species of Dichanthiwm, Iseilema 
and Astrebla occupy comparatively smaller areas between the woodlands, and 
the southern area is dominated by Triodia grassland and open Acacia shrubland. 


The poorly-drained black soils of the Barkly Tableland carry extensive 
erassland mainly of species of Astrebla, but including Iseilema, Dichanthium, 
Sorghum and Dactyloctenium with associated forbs. Relatively small areas are 
dominated by small trees and open shrubland, including Hucalyptus pruinosa, 
Ventilago viminalis, Atalaya hemiglauca and Carissa lanceolata. Areas of Triodia 
pungens are features of the southern part of the Tableland, while Acacia cambagei 
shrubland occurs in the south-eastern part. 


In the Central Australia District, various Acacia shrublands cover desert 
sandplains, desert loams and desert sandhills. Acacia aneura and A. georginae 
are the most common shrub or small tree species, and there are many associated 
grasses and forbs. Triodia and Plectrachne species dominate the deep sandy areas. 
Open low shrubland and sparse grassland occur in the southern area, including 
Many species in the family Chenopodiaceae. The mountain ranges contain relic 
areas with micro-habitats in which Macrozamia macdonnellui, Iivistona mariae, 
several fern species, and some other unique species occur. 


FLORA 
Two new nomenclatural combinations necessary for this check list are : 


Borreria involucrata (O. Schwarz) Chippendale comb. nov. Basionym 
Spermacoce involucrata O. Schwarz in Fedde Repertorium XXIV 
(1927), 102. 

Canthium schultzuw (O. Schwarz) Chippendale comb. nov. Basionym Plectronia 
schultzii_ O. Schwarz in Fedde Repertorium XXIV (1927), 101. 


TABLE | 
Summary of Numbers of Plant Species in the Northern Territory 


Total Species Percentage 
I Endemism 
District Naturalized Restricted Native 
Native or Native Species 
Adventive Species Only 
C.A. (Central Australia) ag! 1,182 54 701 59 
B.T. (Barkly Tableland) ap 494 8 42 8 
V.R.D. (Victoria River District) 918 9 144 16 
D.G. (Darwin and Gulf District) 1,566 64 895 57 
C.A.-B.T. a. i Ae 1,357 59 794 58 
C.A.-V.R.D. Di: se si: 1,732 61 916 53 
B.T.-V.R.D. sy se Be 1,059 15 210 20 
D.G.-B.T. sii oe be 1,779 66 963 54 
D.G.-V.R.D 1,902 65 1,338 70 


Table 1 shows a summary of the species distribution in the Northern 
Territory. The Darwin and Gulf District is the richest in number of species, 
and this is also the area where many more species should yet be recorded. 
Collecting of specimens during the ‘‘ wet ”’ season, when the forbs are available, 


PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 209 


NORTHERN TERRITORY OF AUSTRALIA 


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G. M. CHIPPENDALE 211 


still presents difficulties owing to the lack of all-weather roads to some areas. 
The trees and shrubs have been reasonably, though not exhaustively, collected. 
Many species in the area need investigation to ascertain not only their taxonomic 
position within the Australian flora, but also their relationship with species in 
neighbouring islands and Asia. 

The Victoria River district has a comparatively low total number of species, 
and it seems likely that more records are yet to be made. The number of species 
in the Barkly Tableland is low, and from experience it seems unlikely that there 
will be any vast increase in this number. Further thorough collecting in both of 
these districts seems necessary. 

Since Chippendale’s (1959) check list for Central Australian plants, the 
number of recorded species has increased from 1,101 to 1,236. Some of this 
increase has been on account of new collections, particularly in the area adjacent 
to Western Australia, but recent revisions of genera have also contributed by 
distinguishing species not previously described. J. H. Willis (personal com- 
munications), from the Melbourne Herbarium, advised of previous misidenti- 
fications and of some undescribed species : 


Styphelia mitchellia F. Muell., recorded from Mt. Sonder, is actually an 
undescribed species of Leucopogon. 

Prostanthera schultzii F. Muell. ex Tate, recorded from Mt. Sonder, is actually 
an undescribed species of Wrixonia. 

EHriostemon argyreus F. Muell. ex Tate, in Central Australia, was a mis- 
identified fragment of a species of Pityrodia from the ‘“‘ Mt. Saunders ” 
area near Coolgardie, Western Australia. 


Goodenia sp. nov. (Palm Valley). 
Goodenia sp. nov. (Mt. Liebig). 
Ricinocarpos sp. nov. (Simpson’s Gap). 
Euphorbia sp. nov. (King’s Canyon). 
Portulaca sp. nov. (King’s Canyon). 


Revisions of many genera in recent years have been most valuable, including 
Pandanus, Cassia, Triodia, Bassia, Enneapogon, Ptilotus, Ficus, Capparis, Sesbania, 
Swainsona, Zornia, Neptunia, Erodium, Drosera, Frankenia, some Hibiscus, 
Gossypium, some Hucalyptus, some Melaleuca, Velleia, Stylidium, Nicotiana, 
Utricularia, Brachycome, Calotis, some Acacia and parts of the families 
Loranthaceae, Sapotaceae, Cruciferae and Acanthaceae. 


Much needed are revisions of the important and extensive Cyperus, 
Fimbristylis, Sida, Eragrostis, Indigofera, Tephrosia, Desmodium, Crotalaria, 
Phyllanthus, Euphorbia, Terminalia, Mitrasacme, Heliotropium, Dicrastylis, 
Neweastlia, Solanum, Eremophila, Borreria, Goodenia, Scaevola, and the family 
Scrophulariaceae. 


Table 1 shows the degree of endemism to be greatest in the Darwin and Gulf 
and the Victoria River Districts, if these adjoining zones are considered as one. 
Central Australia District, and the Darwin and Gulf District show only a slight 
difference in percentage endemism, but considering the probability of unrecorded 
species in the more northern seasonally-wet areas, it seems likely that the Darwin 
Gulf District will ultimately show the greatest percentage of endemism. There 
appears to be little endemism in the Barkly Tableland and Victoria River District. 


In Table 2 it can be seen that ferns are notably more prevalent in the Darwin 
and Gulf District, with a much smaller number in Centra] Australia, and only a 
few species elsewhere. This same pattern is evident in Cyperaceae, Loranthaceae, 
Euphorbiaceae, Myrtaceae, Scrophulariaceae, and Rubiaceae. A similar pattern, 
but with an increased number of species in Central Australia, is shown for 


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212 NORTHERN TERRITORY PLANTS 


Gramineae, Papilionaceae and Caesalpiniaceae. There are approximately equal 
numbers of species in central and northern regions in Liliaceae, Proteaceae, 
Amaranthaceae, Atzoaceae, Mimosaceae, Malvaceae, Sapindaceae, Sterculiaceae 
and Boraginaceae. The number of species in the Central Australia District is 
significantly greater in Portulacaceae, Cruciferae, Zygophyllaceae, Frankeniaceae, 
Solanaceae, Myoporaceae, Goodeniaceae and Compositae. The northern areas 
have a greater number of species in Palmae, Araceae, Hriocaulaceae, Haemodoraceae, 
Orchidaceae, Moraceae, Rutaceae, Tiliaceae, Combretaceae, Loganiaceae, Lenti- 
bulariaceae and Stylidiaceae. 


Table 2 also shows the distribution geographically and botanically, of 
naturalized or adventive species. Few of these are widespread. The majority 
are within the influences of the larger population centres, particularly Darwin 
and Alice Springs. In the check list, Verbena officinalis is recorded by Ewart 
and Davies (1917) and V. macrostachya is recorded by Tate (1896), and supporting 
specimens are in the National Herbarium of Victoria, Melbourne. These two 
taxa have been regarded as native Australian species by Bentham (l.c.). However, 
there has been no further record of these species in the Northern Territory. 


ACKNOWLEDGEMENTS 


I acknowledge gratefully the vast assistance I had in the collection and 
caring for specimens and in the beginning of the compiling of the check list from 
Mr. D. J. Nelson of the Primary Industry Branch, Northern Territory Adminis- 
tration, Alice Springs, and Mr. R. F. Swinbourne, who is now at the Botanic 
Garden, Adelaide. 


Mr. J. Maconochie, Primary Industry Branch, Northern Territory Adminis- 

tration, Alice Springs, assisted in checking some specimens. Mr. J. Willis, 
National Herbarium, Melbourne, advised regarding re-identifications of some 
rare specimens, and Mr. G. Stocker, Forestry and Timber Bureau, Darwin, 
supplied a list and specimens of Melville Island plants. Dr. Paul Fryxell, 
United States Department of Agriculture, Texas, advised regarding four species 
of Hibiscus which are to be described by Dr. F. D. Wilson and Mr. N. Byrnes. 
Advice on other special groups of plants was received from the following : 
Mr. D. Symon, Waite Agricultural Research Institute, Adelaide (Solanum spp.). 
Mr. L. Pedley, Queensland Herbarium, Brisbane (certain unnamed Acacia spp.). 
Mr. M. Lazarides, Division of Land Research, C.S.I.R.O., Canberra (Gramineae). 
Dr. Mary Tindale, National Herbarium of New South Wales, Sydney 
(Pteridophyta). 

The manuscript was examined by the botanists at the Royal Botanic Gardens, 
Sydney, who kindly made a number of suggestions for inclusions and exclusions. 


Mr. K. Lyon, Forestry and Timber Bureau, Canberra, prepared the map in 
ome 


Bibliography 

References quoted in the text are included in the following list, which also has many references 

which support specimens or taxonomic details in the check list. 

ApEMA, F., 1969.—Identities of the herbaceous species of Polygala represented in the Brisbane 
Herbarium. Proc. Roy. Soc. Q., 80: 125-30. 

Arry-SHaw, H. K., 1934.—The genus Dentella in Australia. Kew Bull., 1934, 290-301. 

Aut, 8S. I., 1969.—Senecio lautus complex in Australia. V. Taxonomic interpretations. Aust. 
J. Bot., 17: 161-76. 

ANoN., 1919.—Review of The Flora of the Northern Territory. J. Bot., 57: 69-71. 

Bartey, F. M., 1899-1905.—The Queensland Flora. Parts 1-6, and dex. Brisbane. 

Baitey, L. H., 1935.—Certain ptyochospermate palms of horticulturists. Gentes Herbarum, 
3: 410-37. 

Bartow, B. A., 1962.—Studies in Australian Loranthaceae. I. Nomenclature and new additions. 
Proc. Linn. Soc. N.S.W., 87: 51-61. 

, 1963.—Studies in Australian Loranthaceae. JII. A revision of the genus Lyszana. 

Proc. Linn. Soc. N.S.W.. 88: 137-50. 


PROCEEDINGS OF THE LINNEAN Society or New SourH Wates Vout. 96, Part 4 


G. M. CHIPPENDALE 213 


, 1966.—A revision of the Loranthaceae of Australia and New Zealand. Aust. J. Bot., 
14: 421-99. 
, 1970.—Viscum flexicaule, a new species from Northern Australia. Contrib. from the 
N.S.W. Nat. Herb., 4: 95. 
Beavte, N. C. W., Evans, O. D., and Carouin, R. C., 1962.—Handbook of the Vascular Plants 
of the Sydney District and Blue Mountains. Armidale. 


BENL., G., 1958.—Beitrag zu einer revision der gattung Ptilotus R.Br. Sonderdruck aus den 
Mettteilungen der Botanischen Staatssammlung Munchen, Heft 19-20: 401-9. 
———-—-, 1959.—Beitrag zu einer revision der gattung Ptilotus R.Br. Sonderdruck aus den 
Meitteilungen der Botanischen Staatssammlung Munchen, 2 Teil, Band IIT, 34-43. 
, 1960.—Beitrag zu einer revision der gattung Ptilotus R.Br. Sonderdruck aus den 
Meirtteilungen der Botanischen Staatssammlung Munchen, 3 Teil, Band IIT, 510-18. 
, 1961.—Zwei neue Ptilotus—Arten. Sonderdruck aus den Meitteilungen der Botanischen 
Staatssammlung Munchen, Band IV, 75-82. 
, 1962.—Beitrag zu einer revision der gattung Péilotus R.Br. Sonderdruck aus den 
Meitteilungen der Botanischen Staatssammlung Munchen, 4 Teil, Band IV, 277-84. 
, 1964.—New taxa of Ptilotus. Trans. Roy. Soc. S. Aust., 88: 53-60. 
, 1965.—Beitrag zu eimer revision der gattung Ptilotus R.Br. Sonderdruck aus den 
Meitteilungen der Botanischen Staatssammlung Munchen, 5 Teil, Band V, 563-69. 
, 1967a.—Beitrag zu eimer revision der gattung Ptilotus R.Br. Sonderdruck aus den 
Meitteilungen der Botanischen Staatssammlung Munchen, 6 Teil, Band VI, 493-504. 
, 19676.—The genus Ptilotus R.Br. Aust. Pl., 4: 109-24. 
BENTHAM, G., 1863-78.—Flora Australiensis. Vols. 1-7. l. Reeve, London. 


Buacr, J. M., 1943—57.—Flora of South Australia. Parts 1-4, 2nd edition. Adelaide. 


BuacKkati, W. E., and Grinve, B. J., 1954-65.—How to Know Western Australian Wildflowers. 
Parts 1-3. Perth. 
BuakeE, 8. T., 1937.—Notes on Australian Cyperaceae. I. Proc. Roy. Soc. Q., 47: 89-94. 
. 1938.—The genus Jseilema in Queensland. Proc. Roy. Soc. Q., 49: 82-94. 
. 1939.—A monograph of the genus Eleocharis in Australia and New Zealand. Proc. 
Roy. Soc. Q., 50: 88-132. 
. 1940a.—Studies on Queensland grasses. I. Proc. Roy. Soc. Q., 51: 169-76. 
, 19405.—Monographic studies in the genus Fimbristylis. Part I. University of Q. 
Paper's, 1 (13): 1-14. 
. 1941a.—Studies on Queensland grasses. II. University of Q. Papers, 1 (18): 1-22. 
, 1941b.—New genera of Australian grasses. University of Q@. Papers, 1 (19): 1-12. 
, 1943.—Critical notes on the Gramineae and Cyperaceae of South Australia with deserip- 
tions of new species. Trans. Roy. Soc. S. Aust., 67: 42-61. 
, 1944.—Monographic studies in the Australian Andropogonae. Part I. University 
of Q. Papers, 2 (3): 1-62. 
————,, 1947.—Notes on Australian Cyperaceae. VII. Proc. Roy. Soc. Q., 58: 35-50. 
-, 1952.—The identification and distribution of some Cyperaceae and Gramineae chiefly 
from Australia. Proc. Roy. Soc. Q., 62: 83-100. 
. 1953.—Botanical contributions of the Northern Australia Regional Stuvey. I. Studies 
on Northern Australian species of Hucalyptus. Aust. J. Bot., 1: 185-352. 
, 1954.—Botanical contributions of the Northern Australia Regional Survey. II. 
Studies on miscellaneous Northern Australian plants. Aust. J. Bot., 2: 99-140. 
. 1961—Two new species of Scleria. Blumea, 11 (1): 219-23. 
, 1963.—New or noteworthy plants, chiefly from Queensland. II. Proc. Roy. Soc. Q., 
73: 61-77. 
, 1964.—A revision of Micraira, a genus of grasses of exceptional structure and distri- 
bution. Proc. Roy. Soc. Q., 74: 45-52. 
, 1968.—A revision of Melaleuca leucadendron and its allies. Contrib. from the Q. Herb., 
No. 1: 1-114. 
. 1969a.—Studies in Cyperaceae. Contrib. from the Q. Herb., No. 8: 1-48. 
, 1969b.—Taxonomic and nomenclatural studies in the Gramineae. No. 1. Proc. 
Roy. Soc. Q., 80: 55-84. 
. 1969c.—Taxonomic and nomenclatural studies in the Gramineae. No. 2. Proc. 
Roy. Soc. Q., 81: 1-26. 
, 1971.—A revision of Plectranthus (Labiatae) in Australasia. Contrib. from the Q. Herb., 


se ol thet Loranthaceae of Australia. Part 1. Proc. Linn. Soc. N.S.W., 
a “Tithe Loranthaceae of Australia. Part 2. Proc. Lryn. Soc. N.S.W., 47: 
melee eae Loranthaceae of Australia. Part 3. Proc. Liryn. Soc. N.S.W., 47: 

622d. the Loranthaceae of Australia. Part 4. Proc. Lryn. Soc. N.S.W., 48: 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutrH Wates, Vou. 96, Part 4 


214 NORTHERN TERRITORY PLANTS 


———-—, 1922e.—The Loranthaceae of Australia. Part 5. Proc. Linn. Soc. N.S.W., 49: 
79-96. 

, 1925.—The Loranthaceae of Australia. Part 6. Proc. Linn. Soc. N.S.W., 50: 1-24. 

, 1965.—A Key to the Hucalypts. 3rd ed. Forestry and Timber Bureau, Canberra. 

Boomsma, C. D., 1964.—A description of a new mallee species of Hucalyptus from central Australia. 
Trans. Roy. Soc. S. Aust., 88: 115-16. 

Borssum WAALKES, J. VAN, 1966.—Malesian Malvaceae revised. Blumea, 14: 1-213. 

BremeKkamp, C. HE. B., 1962.—Remarks on the position of some Australian Acanthaceae. Acta 
Botanica Neerlandica, 11: 195-200. 

, 1964.—On the systematic position of the Australian Nelsonias and Thunbergias and 
of the Ruellia species which by Domin were referred to Aporuellia Clarke. Koninkl. Nedert. 
Akademie van Wetenschaffen Amsterdam, Proc. Series C 67 (5): 301-6. 

Brenan, J. P. M., and Brummirr, R. K., 1965.—The variation of Dichrostachys cinerea (1..) 
Wight and Arn. SBoletim da Sociedade Broteriana, 29: 61-115. 


Brraes, Barsara G., and Jonnson, L. A. 8., 1968.—The status and relationships of the 
Australasian species of Typha. Contrib. from the N.S.W. Nat. Herb., 4: 57-69. 
BorBipGeE, Nancy T., 1940—-1.—A revision of the Australian species of Hnneapogon Desv. Proc. 
Linn. Soc. of London, 153rd session (1): 1-91. 
, 1953.—The genus Triodia R.Br. Aust. J. Bot., 1: 121-84. 
————, 1958.—A monographic study of Helichrysum subgenus Ozothamnus (Compositae) and 
of two related genera formerly included therein. Aust. J. Bot., 6: 229-84. 
, 1960.—Further notes on Triodia R.Br. with descriptions of five new species and one 
variety. Aust. J. Bot., 8: 381-94. 
, 1965.—The Australian species of Sesbania Scopoli (Leguminosae). Aust. J. Bot., 
13: 103-41. 
Byrnes, N., 1971.—New species of Lophopetalum (Celastraceae), a genus new for Australia. Proc. 
Linn. Soc. N.S.W. In press. 
Carouin, R. C., 1958.—The species of the genus Hrodiwm L’Her. endemic to Australia. Proc. 
Linn. Soc. N.S.W., 83: 92-100. 
——, 1967.—The genus Velleia Sm. Proc. Linn. Soc. N.S.W., 92: 27-57. 
CHIPPENDALE, G. M., 1959.—Check list of central Australian plants. Trans. Roy. Soc. S. Aust., 
82: 321-38. 
, 1960.—Contributions to the flora of central Australia. No. 1. Trans. Roy. Sec. 
S. Aust., 83: 199-203. 
—, 1961.—Contributions to the flora of central Australia. No. 2. Trans. Roy. Soc. 
S. Aust., 84: 99-103. 
, 1963.—Contributions to the flora of central Australia. No. 3. Trans. Roy. Soc. 
S. Aust., 86: 7-9. 
, 1964.—Records of Northern Territory plants. Trans. Roy. Soc. S. Aust., 88: 175-80. 
CuemeEnt, I. D., 1957.—Studies in Sida. I. Contrib. from the Gray Herb. of Harvard Univ., 180 : 
3-91. 
CocHRANE, G. R., 1967.—The description and mapping of vegetation in Australia. Vic. Nat., 
84: 299-316. 
Cooper, R. C., 1956.—The Australian and New Zealand species of Pittosporum. Annals of the 
Missouri Bot. Gard., 43: 87-188. 
CopPpELAND, KH. B., 1947.—Genera Filicum. The Ronald Press Co., New York. 


Corner, KE. J. H., 1960.—Taxonomie notes on Ficus L., Asia and Australasia, Sections 1—4. 
The Gard. Bull., Singapore, 17 (3): 368-485. 

—, 1965.—Check List of Ficus in Asia and Australasia, with keys to identification. The 
Gard. Bull., Singapore, 21: 1-186. 

Curtis, WINIFRED M., 1956—67.—The Student’s Flora of Tasmania. Parts 1-3. Govt. Printer, 
Tasmania. 

Davis, Gwenpa L., 1948.—Revision of the genus Brachycome Cass. Part 1. Australian species. 
Proc. Linn. Soc. N.S.W., 73: 142-241. 

————, 1949.—Revision of the genus Brachycome Cass. Part 3. Description of three new 
Australian species and some new locality records. Proc. Linn. Soc. N.S.W.. 74: 145-42. 

————, 1952.—Revision of the genus Calotis R.Br. Proc. Linn. Soc. N.S.W., 77: 146-88. 

————, 1955.—Supplementary notes on the genus Brachycome Cass. Description of five new 

Australian species and some new locality records. Proc. Linn. Soc. N.S.W.. 79: 203-10. 

——, 1956.—Revision of the genus Podolepis Labill. Proc. Linn. Soc. N.S.W., 81: 245-86. 

DeListe, D. G., 1963.—Taxonomy and distribution of the genus Cenchrus. Iowa State J. Sc., 
37 (3): 259-351. 

Etcuuer, Hs., 1965.—Supplement to J. M. Black’s Flora of S. Aust. S. Aust. Branch of the British 
Se. Guild Handbook Committee, Adelaide. 

Erickson, Rica, 1958.—Triggerplants. Paterson Brokensha Pty. Ltd., Perth. 

————, 1968.—Plants of Prey in Australia. Lamb Publications Pty. Ltd., Western Australia. 

————, and Wiis, J. H., 1966.—Some additions to Australian Stylidiaceae. Vic. Nat., 
83: 107-12. 


PROCEEDINGS Or THE LINNEAN Society or New SoutH Waters, Vou. 96, Part 4 


G. M. CHIPPENDALE 215 


Ewarr, A. J.. and Daviss, Otrve B., 1917.—The Flora of the Northern Territory. Minister for 
Home and Territories, Melbourne. 
Ewart, A. J., and JARRETT, PHytiis H., 1927.—\Contribution to the flora of Australia. No. 33. 
Additions to the flora of the Northern Territory and locality records. Proc. Roy. Soc. Vic., 
39: 154-61. 
, 1928.—Contribution to the flora of Australia. No. 34. Additions to the flora of the 
Northern Territory and locality records. Proc. Roy. Soc. Vic., 40: 83-87. 
Forpg, N., and Isrne, E. H., 1958.—Acacia calcicola, a new species of importance to the ecology 
of the Australian arid zone. Trans. Roy. Soc. S. Aust., 81: 153-60. 
FRYXELL, P. A., 1965.—A revision of the Australian species of Gossypium with observations on the 
occurrence of Thespesia in Australia. Aust. J. Bot., 13: 71-102. 
, 1968.—A redefinition of the tribe Gossypieae. Bot. Gaz., 129: 296-308. 
, 1969.—A classification of Gossypium L. (Malvaceae). Taxon, 18: 585-91. 
GARDNER, C. A.. 1952.—Flora of Western Australia. Vol. 1, Gramineae. Government Printer, 
Perth. 
HarrLey, W., 1942.—New species and varieties of Hriachne R.Br. J. Linn. Soc. Botany, 52 (344) : 
342-48. 
HeEnxKER, H., 1967.—Die gattung Szgesbeckia. Arch. Nat. Meckl., X1: 7. 
HusBarp, C. H., 1928.—The genus Astrebla, or Mitchell grasses. Bull. Misc. Inf., No.7: 2 
, 1935.—Species novae vel minus cognitae. Hook. Ic. Pl., tt. 3283-6. 
————, 1939.—Species minus cognitae. Hook. Ic. Pl., t. 3385. 
, 1952.—Whiteochloa, a new genus of grasses from the Northern Territory of Australia. 
Proc. Roy. Soc. Q., 62: 109-12. 
Hucues, D. K., 1923.—The genus Panicum of the Flora Australiensis. Bull. Misc. Inf. Royal 
Botanic Gardens, Kew, 9: 305-32. 
Isine, E. H., 1964.—The species of Bassia All. in Austraha. Trans. Roy. Soc. S. Aust., 88 : 63-110. 
Jacoss, M., 1964.—The genus Capparis from the Indus to the Pacific. Blumea, 12: 385-541. 
JANSEN, P., 1930.—Two new species of Hriachne. Mededeelingen’s Rijks Herbarium, Leiden, 
No. 59: 1-2. 
JEDWABNICK, ELISABETH, 1924.—Eragrostidis speciarum imprimus ad herb. Berol. Hamburg, 
Mona., Regimont digestarum compactus. Bot. Archiv., V. Band: 177-216. 
Jounson, L. A. S., 1959.—The families of Cycads and the Zamiaceae of Australia. Proc. Linn. 
Soc. N.S.W., 84: 64-117. 
KostERMANS, A. J. G. H., 1965.—A monograph of the genera Maranthes Bl]. and Cyclandrophora 
Hassk. of the Asiatic and Pacific area. Candollea, 20: 103-42. 
, 1969.—New and critical Malesian plants. VIII. Retnwardtia, 7: 443-49. 
LAZARIDES, M., 1959.—The Australian species referred to Ichnanthus Beauv. Aust. J. Bot.. 
7: 328-46. : 
, 1970.—The Grasses of Central Australia. Australian University Press, Canberra. 
Lez, Atma T., 1948.—The genus Swainsona. Contrib. from the N.S.W. Nat. Herb., 1: 131-271. 
————, 1962.—Notes on Lomandrain N.S.W. Contrib. from the N.S.W. Nat. Herb., 3: 151-64. 
LEeENHouTs, P. W., 1959.—A monograph of the genus Canarium. Blumea, 9: 275-475. 
MouHLENBROCK, R. H., 1961.—A monograph of the leguminous genus Zornia. Webbia, 16: 1-141. 
Morton, J. K., 1966.—A revision of the genus Aneilema R.Br. J. Linn. Soc. (Bot.), 59: 431-78. 
Monk, W. J. pe, 1962.—Preliminary revision of some genera of Malaysian Papilionaceae. III. A 
census of the genus Crotalaria. Reinwardtia, 6: 195-223. 
Nean, P. T., 1965.—A revision of the genus Wrightia (Apocynaceae). Annals Missouri Bot. 
Gard., 52: 114-75. 
Oostsrroom, 8S. J., 1934.—A monograph of the genus Hvolvulus. Mededeelingen van het Botanical 
Museum en Herbarium van de Rijksuniversiteit te Utrecht, 14: 1-267. 
, 1938.—The Convolvulaceae of Malaysia. 1. Blumea, 3: 62-94. 
, 1939.—The Convolvulaceae of Malaysia. 2. Blumea, 3: 267-371. 
—————, 1940.—The Convolvulaceae of Malaysia. 3. Blumea, 3: 481-582. 
, 1943.—The Convolvulaceae of Malaysia. 4. Blumea, 5: 339-411. 
, 1945.—The Convolvulaceae of Malaysia. 5. Blumea, 5: 689-91. 
Payvens, J. P. D. W., 1957.—Revision of some Papuan Liliaceae. Nova Guinea, n.s. 8 (2): 390. 
————, 1967.—A monograph of the genus Barringtonia (Lecythidaceae). Blumea, 15: 157-263. 
PEpDLEY, L., 1964a.—Notes on Acacia, chiefly from Queensland. I. Proc. Roy. Soc.Q., 74: 53-59. 
, 1964b.—Notes on Acacia, chiefly from Queensland. II. Proc. Roy. Soc. Q., 75: 


Or 
in 
fon) 
~I 


29-35. 
, 1969.—Notes on Acacia, chiefly from Queensland. III. Contrib. from the Q. Herb., 
No. 4: 1-7. 


Puitcox, D., 1968.—Revision of the Malesian species of Lindernia All. Kew Bull., 22: 1-72. 

PotHity, R. M., 1968.—Miscellaneous notes on African species of Crotalaria L. II. Kew Buill., 
22 (2): 169-348. ve 

Ranperia, A. J., 1960.—The composite genus Blumea, a taxonomic revision. Blumea, 10: 


176-317. 
Raven, P. H., 1963.—The Old World species of Ludwigia (including Jussiaea) with a synopsis of 


the genus. Reinwardtia, 6: 327-427. 


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216 NORTHERN TERRITORY PLANTS 


Rosyns, A., 1963.—Essai de monographie du genre Bombaz. Bull. Jard. Bot. Bruax., 33 (1): 
1-316. 

Royen, P. van, 1957.—Revision of the Sapotaceae of the Malaysian area in a wider sense. VII. 
Planchonella Pierre. Blumea, 8: 235-445. 

Sauer, J., 1964.—Revision of Canavalia. Brittonia, 16 (2): 106-81. 

ScHoppE, R., 1963.—A taxonomic revision of the genus Millotia Cassini (Compositae). Trans. 

' Roy. Soc. S. Aust., 87: 209-41. 

ScHwakRz, O., 1927.—Plantae novae vel minus cognitae Australiae tropicae. Meddes Repertorium, 
14: 80-109. 
Suaw, ExizaBetu A., 1965.—Taxonomic revision of some Australian endemic genera of Cruciferae. 
Trans. Roy. Soc. S. Aust., 89: 145-258. 
SmirH, L. S., 1956.—New species of and notes on Queensland plants. Proc. Roy. Soc. Q., 67: 
29-40. 
Sprcur, R. L., 1958a.—The Gymnospermae and Angiospermae collected on the Arnhem Land 
Expedition. The American-Australian Scientific Hupedition to Arnhem Land. Vol. 3, Botany 
and Plant Ecology, No. 9: 185-317. 
STauFFER, H. U., 1959.—Revisio Anthobolearum. Mitteclungen aus dem Botanischen Museum 
der Universitat Zurich, 213: 1-260. 
Sreents, C. C. G. J. van, 1929.—A revision of the Queensland Bignoniaceae. Proc. Roy. Soc. Q., 
41: 39-58. 
Sr. Joun, H., 1962.—Revision of the genus Pandanus Stickman. Part 13. Pandanus in the 
Northern Territory, Australia. Pacific Sc., 16: 409-28. 
SuMMERHAYES, V. S., 1929.—A revision of the Australian species of Frankenia. J. Linn. Soc. 
Bot., 48: 337-87. 
Symon, D. E., 1966.—A revision of the genus Cassia L., Caesalpiniaceae in Australia. Trans. 
Roy. Soc. S. Aust., 90: 73-146. 
TiInDALE, Mary D., 1958.—The Pteridophyta of Arnhem Land. The American-Australian Scientific 
Hapedition to Arnhem Land. Vol. 3, Botany and Plant Hcology, No. 9: 171-84. 
, 1968.—Notes on Australian taxa of Acacia. No. 1. Contrib. from the N.S.W. Nat. 
Herb., 4: 73-78. 
, 1970.—Notes on Australian taxa of Acacia. No. 2. Contrib. from the N.S.W. Nat. 
Herb., 4: 137-43. 
Waits, C. T., 1936.—Contribution to the Queensland Flora, No. 5. Proc. Roy. Soc.Q., 47: 1-35. 
, 1950.—Additions to the flora of Arnhem Land. Proc. Roy. Soc. Q., 61: 55-58. 
Wits, J., 1942.—Botanical bulls. Vic. Nat., 59: 72. 
, 1945.—More botanical bulls. Vic. Nat., 61: 175. 
, 1962.—A Handbook to Plants in Victoria. Vol. 1. Melbourne University Press, 
Melbourne. 
, 1967.—Systematie notes on the indigenous Australian flora. Muelleria, 1: 117-63. 
—, 1969.—Prostanthera schultzii seen in flower for the first time. Vic. Nat., 86: 132. 
Witson, F. D., and Byrnes, N.—Two new species of Hibiscus, Section Furcaria (Malvaceae) 
from Australia. Proc. Linn. Soc. N.S.W., 95: 194-97. 

Witson, P. G., 1960.—A consideration of the species previously included within Helipterum 
albicans (A. Cunn.) DC. Trans. Roy. Soc. S. Aust., 83: 163-77. 

Winpvter, D. R., 1966.—A revision of the genus Neptunia (Leguminosae). Aust. J. Bot., 14: 
379-420. 

Wirt, H. C. D. pz, 1956.—A revision of Malaysian Bauhinieae. Reinwardtia, 3: 381-541. 


CHECK EIST 


* Naturalized. 
+ Adventive. 
C.A. IBA, =. \WelolD, IDs @e Gi 


LYCOPODIACEAE 
Lycopodium cernuum lL. .. a as ae se af we x x 
SELAGINELLACEAE 
Selaginella ciliaris (Retz.) Spring ay at ate aa ss Sis Xx 
Selaginella uliginosa (Labill.) Spring .. vie e. ies He o6 x 
JSOETACEAE 
Tsoetes humilior F. Muell. ex A.Br. se a, ae x< 
PSILOTACEAE 
Psilotum nudum (L.) Griseb. te “At, Fe fe x 


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G. M. CHIPPENDALE 2NF 


GAL) SB ya eee: 


OPHIOGLOSSACEAE 
Helminthostachys zeylanica (Li.) Hook. .. Se ‘ic ic sar a x 
Ophioglossum lusitanicum L. ssp. coriaceuwm (A. Cunn.) 

Clausen .. = a aie rp x 
Ophioglossum mulgatum ee 


x: 


SCHIZAEACEAE 
Lygodium flexuosum (1.) Swartz .. 
Lygodium japonicum Swartz 
Lygodium microphyllum (Cav.) R. Br. 
Schizaea bifida Willd. 
Schizaea dichotoma (L.) Sm. 


XX XX X 


GLEICHENIACEAE 
Dicranopteris linearis (Burm. f.) Underwood .. “igs ae oo o6 x 
Dicranopteris sp... bk ae ae 5 on ae Sc x x 
Gleichenia sp. aA a be ae a a ee O00 x 


DENNSTAEDTIACEAE 
Histiopteris incisa (Thunb.) J.Sm. ae ee Ki x 


HyMENOPHYLLACEAE 
Selenodesmium sp. .. ot ists a ae 6.0 dre as 5c x 


LINDSAEACEAE 
Lindsaea ensifolia Swartz .. 6 Ps a0 ae x o¢ 50 x 
Lindsaea frasert Hook. on oi ae a O¢ ate Oc re x 
Lindsaea orbiculata Mett. x 


ADIANTACEAE 
Adiantum hispidulum Swartz oot ai a ae x 
Adiantum philippense L. .. be x 
Cheilanthes distans (R.Br.) Mett. Ey an fas ae Ke x 
Cheilanthes fragillima F.Muell. .. | Dit 50 2c o-6 De oe x 
Cheilanthes javensis Moore im a ee eo om x 
Cheilanthes lasiophylla Pic.-Ser. .. oe aa a x 
Cheilanthes paucijuga Benth. ae ae Bie so ae aa x 
Cheilanthes ? sciadioides Domin .. Ms a Se o is x 
Cheilanthes siebert Kuntze ae ae se ae 
Cheilanthes tenuifolia (Burm. f.) Swartz var. tenuzfolia. . 
Cheilanthes tenuifolia (Burm. f.) Swartz var. nudiuscula 
(R.Br.) F. M. Bail. Rh: ae ae 5 ote Ss 
Cheilanthes vellea (R.Br.) F. Muell. << Ae as xx Ls x 
Gymnogramma reynoldsti (F. Muell.) J. M. Black ahs x ae Me 
Neurosoria pteroides (R.Br.) Mett. ae ed ‘ ig a Ss 
Platyzoma microphyllum R.Br... is 50 00 oe ffs 20 x 
Taenitis blechnoides (Willd.) Swartz as iG ne vo a Se 


x xX 
M6 
x 


PTERIDACEAE 
Acrostichum speciosum Willd. ie 0% ois ar aie is Be x 
Pteris tremula R.Br. ae sig te =e & x 


POLYPODIACEAE 
Drynaria quercifolia (L.) J.Sm. ee 
Microsorium scolopendria (Burm. f.) Copel. 


xX 


PARKERIACEAE 
Ceratopteris thalictroides (L.) Brongn. .. ae a Bie ae oe x 


DAVALLIACEAE 
Nephrolepis cordifolia (L.) Pre Ce a i hs x He Pr we 
Nephrolepis obliterata (R.Br.) C fe ec ae 53 a fe x 


ASPLENIACEAE 
Pleurosorus rutifolius (R.Br.) Fée ; ar ~< 
Pleurosorus subglandulosus (Hook. et Grev. \M . Tindale x 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VoL. 96, Part 4 


218 NORTHERN TERRITORY PLANTS 
C.A. B.T V.R.D. D. & G. 
THELYPTERIDACEAE 
Cyclosorus gongylodes (Schkuhr) Link .. Bs ai x x 
Cyclosorus sp. we ee cies : x 
BLECHNACEAE 
Blechnum indicum Burm. f. x 
Blechnum orientale L. wt se ake x 
Stenochlaena palustris (Burm. f.) Bedd. .. x 
ASPIDIACEAE 
Polystichum proliferum (R.Br.) Presl  .. se a x 
MARSILEACEAE 
Marsilea angustifolia R.Br. bie ee mK ee x « 
Marsilea crenata Presl i a Ny. ke ay x x 
Marsilea drummondw A.Br. ae Ny ne a x ie me x 
Marsilea exarata A.Br. Bd ae fe ne ae x x x 
Marsilea hirsuta R.Br. or ve i a: ae x x ae 
Marsilea mutica Mett. bar ate ¥ ay ae 4 x x 
AZOLLACEAE 
Azolla pinnata R.Br. x 
CyYCADACEAE 
Cycas media R.Br. x 
Cycas sp. x 
ZAMIACEAE 
Macrozamia macdonnelli (F. Muell. ex Mig.) A.DC. .. ~ 
CUPRESSACEAE 
Callitris glauca R.Br. ex R.T.Bak. et H.G.Sm. Se: < ae ee 
Callitris intratropica R. T. Bak. et H.G.Sm. x x 
TYPHACEAE 
Typha domingensis Pers. i BS ms ae x x 
PANDANACEAE 
Pandanus arnhemicus St.John. x 
Pandanus basedowiu C. H. Wright x 
Pandanus convexus St. John x“ 
Pandanus darwinensis St. John 30 x 
Pandanus integer St. John x x 
Pandanus latifructus St. John x 
Pandanus semiarmatus St. John x 
Pandanus spechtii St. John x 
Pandanus thermalis St. John ; x 
Pandanus yirrkalaensis St. John .. x 
POTAMOGETONACEAE 
Potamogeton tenuicaulis F. Muell.. . Me ac ae x 
Potamogeton tricarinatus A. Bennett ie se is Sg 
Ruppia maritima L... om a wn a ote x 
NALIADACEAE 
Naias marina L. Ne 
APONOGETONACEAE 
Aponogeton elongatus (F. Muell.) Benth. x 
ScHEUCHZERIACEAE 
Triglochin calcitrapa Hook. x 
Triglochin centrocarpa Hook. as S 
Triglochin hexagona J.M. Black .. sie ae x 5 a 
Triglochin procera R.Br. var. dubia (R.Br.) Benth. x x 
Triglochin procera R.Br. var. procera x x 
PROCEEDINGS OF THE LINNEAN Socrety or New SourH Wates, Vow. 96, Part 4 


G. M. CHIPPENDALE 219 


C.A. Bai V.RD. D. & G. 


ALISMATACEAE 
Alisma acanthocarpum F.Muell. .. SE a 52 is te x Me 
Alisma oligococcum F. Muell. 54 oe i ae is ae x Ke 


HyYDROCHARITACEAE 
Blyxa octandra (Roxb.) Planch. ex Thw. 
Blyxa sp. 
Halophila ovalis (R. ‘Br. ) Hook. f. 
Hydrilla verticillata (L.f.) Royle 
Ottelia alismoides Pers. is at ae me ne ve 
Ottelia ovalifolia (R.Br.) L. C. Rich a ae 9: xe as ay Sy 
Vallisneria spiralis L. Ses us us a uf ee Ai Ate x 


x XX XX 


GRAMINEAE 

Acrachne racemosa (Heyne ex R. et 8.) Ohwi 
Alloteropsis semialata (R.Br.) Hitche. ; 
Amphipogon caricinus F. Muell. var. caricinus 
Amphipogon caricinus F. Muell. var. sericeus J. Vickery 
Aristida anthozxanthoides (Domin) Henr. : = 
Aristida biglandulosa J. M. Black 

Aristida browniana Henr. .. 5 

Aristida capillifolia Henr. 

Aristida contorta F. Muell... 

Aristida echinata Henr. . 

Aristida euserta S. T. Blake 

Aristida hygrometrica R.Br. Be Be - 23 
Aristida inaequiglumis Domin .. iG ae Ee x oe x 
Aristida ingrata Domin .. ie = e Pe ae _ ee x 
Aristida jerichoensis Domin ae Be mA ae 
Aristida latifolia Domin .. as wt uf se x 
Aristida macroclada Henr. ae 56 ue ae it is ie x 
Aristida nitidula (Henr.) 8. T. Blake ex J. M. Black .. x 

Aristida obscura Henr. ae oo as Si ae x : ue 

Aristida pruinosa Domina .. Pe se a a x x x x 
Aristida queenslandica Henr. aie tae aE ae ae sf EN x 
Aristida schultz Mez ae 2g ae a Ss 
Aristida strigosa (Henr.) S. T. Blake Re ae Be x 

Arundinella nepalensis Trin. A : : 
Arundinella setosa Trin. .. 

Astrebla elymoides F. Muell. ex F. M. Bail. ae s< 
Astrebla lappacea (Lindl.) Domin Ne x 
Astrebla pectinata (Lindl.) F. Muell. ex Benth. x 
Astrebla squarrosa C. E. Hubbard ; atc pe Be 
+Avena fatua Iu. 4 ie +4 ae x “4 
Bambusa arnhemica F. Muell. is ae ae ae 2 3 sf x 
Rothriochloa decipiens (Hack.) ©. E. Hubbard var. 

cloncurrensis (Domin) C. E. Hubbard th ae is x 

Bothriochloa ewartiana (Domin) C. E. Hubbard St x ae = 
Bothriochloa bladhi (Retz.) S.T. Blake .. a Ss x x x SK 
Brachiaria argentea (R.Br.) Hughes st ie ae Se oe =e 4 
*Brachiaria distachya (1..) Stapf x 
Brachiaria gilesii (Benth.) Chase a od nes x a = 
Brachiaria holosericea (R.Br.) Hughes .. Me i x as se 
Brachiaria miliiformis (C. B. Presl) Chase x : 
Brachiaria mutica (Forsk.) Stapf we es ag : 

Brachiaria notochthona (Domin) Stapf .. Be = x ev se 
Brachiaria piligera (F. Muell. ex Benth.) Hughes ae x x< x x 
Brachiaria polyphylla (R.Br.) Hughes .. : a sf: _. o. Sx 
Brachiaria praetervisa (Domin) C. E. Hubbard | x 3 es oi: 
Brachiaria ramosa (L.) Stapf var. grandiflora Hughes | = Be x Xx 
Brachiaria reptans (l.) C. A. Gardn. et C. E. Hubbard me as x x 
Brachyachne ciliaris (Benth.) C. E. Hubbard .. s S< =e é 
Brachyachne convergens (F. Muell.) Stapf ae a Ss x 
Brachyachne tenella (R. Br.) C. E. Hubbard ae his oh 6 x 
Capillipedium parviflorum (R.Br.) Stapf a = ot se e x 
Capillipedium spicigerum : T. Blake  .. ic oc sé i ~ XS 


XXX: 
XX X XX 


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PROCEEDINGS OF THE LINNEAN SocrETY or New SoutH Wates, Vor. 96, Part 4 


220 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. & G. 


*Cenchrus brown R. et S.. . * ae Be 50 de ae 
*Oenchrus ciliaris L.. 5s se as ae mY x >< ~< 
*Cenchrus echinatus The as AS a 8G << ae Li, 

Cenchrus elymoides F. Muell. are Be ie “i ae Pe x 


*Cenchrus setigerus Vahl 

*Cenchrus tribuloides L. : 

Chamaeraphis hordacea R.Br. kg ys ae a8 
Chionachne cyathopoda (F. Muell.) ‘F. Muell. ex Benth... # a x 
Chionachne hubbardiana Henr. .. ue a, x F 
Chloris acicularis Lindl. .. ae fa ue we x 

*Chloris barbata (L.) Swartz : ae os be Bae a 

Chloris pectinata Benth. .. Be ots ae ee x x 

Chloris pumilio R.Br. : 
Chloris scariosa F.Muell. .. 
Chloris virgata Swartz ; 
Chrysopogon fallax 8. T. Blake 
Chrysopogon latifolius 8. T. Blake : =i BY: a ae 
Chrysopogon pallidus (R.Br.) Trin. ex Steud. .. we x x 
Chrysopogon setifolius Stapf ea ae he i 

Coelorhachis rottboellioides (R.Br.) A. Camus 
Cymbopogon ambiguus A. Camus ie 
Cymbopogon bombycinus (R.Br.) Domin 
Cymbopogon obtectus 8. T. Blake .. Pe 
Cymbopogon procerus (R.Br.) Domin 
Cymbopogon refractus (R.Br.) A. Camus 
*Cynodon arcuatus J.S. Pres] ex C. B. Presl os oe 
*Cynodon dactylon (L.) Pers. ae ae Ae ae x =: 
*Dactyloctentum aegyptium (L.) Beauv. .. an ae ae x 
Dactyloctenitum radulans (R.Br.) Beauv. 
Danthonia bipartita F. Muell. 

Dichanthium affine (R.Br.) A. Camus 
Dichanthium annulatum (Forsk.) Stapf .. 
Dichanthium fecundum §S. T. Blake 
Dichanthium sericeum (R.Br.) A. Camus 
Dichanthium tenuiculum (Steud.) 8. T. Blake 
Digitaria adscendens (H.B.K.) Henr. . 
Digitaria ammophila (F. Muell.) Hughes 
Digitaria brown (R. et S.) Hughes 
Digitaria coenicola (F. Muell.) Hughes 
Digitaria ctenantha (F. Muell.) roe ae 
Digitaria eriolepis Henr. sie SG Sct nig 
Digitaria gibbosa (R.Br.) Beauv. ae ees = me eat x 
Digitaria leucostachya (Domin) Henr. 
Digitaria longiflora (Retz.) Pers. 
Digitaria papposa (R.Br.) Beauv. 
Digitaria robusta Hughes .. 
Digitaria sanguinalis (L.) SestDe Be oe RD Si: ht ah Ed 
Digitaria stenostachya (Benth.) Hughes .. x ore ae a x 
Dimeria ornithopoda Trin. au aie : 

Diplachne fusca (L.) P. Beauv. 
Diplachne muellert Benta. Bes 
Dinlachne parviflora (R.Br.) Benth. 
Echinochloa colonum (L.) Link x ae se 
Echinochloa crus-galli (L.) Beauv. ee a a ar so S< 
Echinochloa turneriana Domin .. on Se og oa x 
Echinochloa stagnina (Retz.) Beauv. 

Ectrosia agrostoides Benth. 

Ectrosia laxa 8. T. Blake a se ee es s a 
Ectrosia leporina R.Br. var. leporina ie i on x 4 x 
Eetrosia leporina R.Br. var. micrantha Benth. a bys Sg 

Ectrosia scabrida ©. E. Hubbard he Be eh ats x as 
Kictrosia schultzii Benth. .. hs ae snes at as es x 
Hleusine indica (L.) Gaertn. : 

Elyonurys citreus (R.Br.) Munro ex Benth. ; an ae Be 
Hlytrophorus spicatus (Willd.) A. Camus oi fe x x x 


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PROCEEDINGS OF THE LINNEAN Soctety or New SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 271 


C.A. Wsiho | (Wolb ID, ID, te (61, 


Enneapogon avenaceus (Lindl.) C. EK. Hubbard .. 

Enneapogon clelandwu N.T. Burbidge 

Enneapogon cylindricus N. T. Burbidge 

Enneapogon glaber N. T. Burbidge 

Enneapogon lindleyanus (Domin) C. E. Hubbard 

Enneapogon oblongus N. T. Burbidge 

Enneapogon pallidus (R.Br.) Beauv. 

Enneapogon planifolius N.T. Burbidge 

Enneapogon polyphyllus (Domin) N. T. Burbidge 

Enneapogon purpurascens (R.Br.) Beauv. 

Eragrostis barreliert Daveau 

Bragrostis bleesert Pilger : 

Eragrostis browni (Kunth.) Nees 

t+EHragrostis cilianensis (All.) Link ex Lutati 

Eragrostis concinna (R.Br.) Steud. 

Eragrostis confertiflora (J. M. Biss) J. M. Black 

Hragrostis cumingw Steud.. 

Hragrostis desertorum Domin 

Hragrostis diels Pilger .. 

Hragrostis elongata (Willd.) J. F. Ji deeds 

Eragrostis eriopoda Benth. 

Eragrostis falcata Gaud. 

Eragrostis tnterrupta Beauv. ae 

Eragrostis japonica (Thunb.) Trin. 

Hragrostis kennedyae F. Turner 

Hragrostis lacunaria F. Muell. ex Benth. 

Eragrostis laniflora Benth. 

Hragrostis leptocarpa Benth. ‘ 

Eragrostis parviflora (R.Br.) Trin. 

Eragrostis schultzii Benth. : 

Eragrostis setifolia Nees : 

Hragrostis speciosa (R. et S.) Steud. : 

Eragrostis stenostachya Steud. var. floribunda Benth. be 

Hragrostis tenax Jedw. a a Be oe as a3 x< x 

Eragrostis xerophila Domin Se, ae eae re Shs x we ax ane 

Eriachne agrostidea F. Muell. a ue Be sh a bs me 3 

Eriachne aristidea F. Muell. one a sa x a x A 

Eriachne armitiw F. Muell. ex Benth. My: a a x Fs Pa x 

Eriachne avenacea R.Br. .. : ae 48 te ee at x S< 

Eriachne basedowit W. Hartley .. Pee ae ae Pe Be x 

Eriachne benthami W. Hartley .. ae whe “i x am : a 

Eriachne burkittu Jansen .. oA a as eee a =f ae x 

Hriachne ciliata R.Br. i are A a ats ai x x x 

Hriachne festucacea F. Muell. are ae a a6 ae eis >< 

Eriachne filiformis W. Hartley Su Ses Pe a bu x 

Hriachne glauca R.Br. ae oe 53 oie oe te x x 

Eriachne helmsii (Domin) W. Hartley 

Hriachne isingiana J. M. Black ae sits oF ae se 

Hriachne laidlawu Jansen ae 5 a a A a ae x 

Eriachne melicacea F. Muell. os 35 Sie a x< x x 

Hriachne mucronata R.Br. var. mucronata ‘ x 

Eriachne mucronata Ji.Br. var. desertorum C. A. Gardn. x 
x 
x 


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Hriachne nervosa Ewart et Cookson 
Eriachne obtusa R.Br. var. obtusa Bi 
Hriachne obtusa R.Br. var. major Ewart et Davies .. ag we oe bx 
Eriachne pulchella Domin .. an oe ae as x a ait oe 
Eriachne rara R.Br. , ae a ve a fe ~ ee x 
EHriachne schultziana F. Muell. ext or oe Be is Ba Hee s 
Eriachne scleranthoides F.Muell. .. aye ae Se >< v3 a 
EHriachne setacea Benth. .. avo Br is 6 $y a we x 
Eriachne squarrosa R.Br. .. oe Mt ae x x 
Hriachne stipacea F. Muell. var. hirsuta W. Hartley Rid ae os ay Xx 
Hriachne sulcata W. Hartley ae : of ae an ay SK ee 
EHriachne triodioides Domin ae a a ue 3 sh a x 
Eriachne triseta Nees et Steud. .. aE Als AF A ioe t x 
Eriochloa australiensis Stapf ex Thell. .. a ae x 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vou. 96, Part 4 


222 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. & G. 


Eriochloa crebra 8. T. Blake : 
Hriochloa procera (Retz.) C. E. Hubbard — : 
Eriochloa pseudoacrotricha Steptoe ex Thell. ) C. E. Hubbard 
ex 8. T. Blake .. : x oe Bs. 
Eulalia fulva (R.Br.) Kuntze sts a as x x x 
Eulalia mackinlayt (F. Muell. ex Benth.) S. T. Blake 
aff. Hulalia (undescribed sp.) : : ae ate 
Heterachne abortiva (R.Br.) Druce ; and ves a wet 
Heteropogon contortus (L.) Beauv. ex R. et S. we Sh x x 
Heteropogon triticeus (R.Br.) Stapf sis a aie 
+Hordeum leporinum Link .. 30 a0 36 x 
Hymenachne acutigluma (Steud.) Gillil. : He 
Imperata cylindrica (L.) Beauv. var. major (Nees) C. E. 
Hubbard a he te af tis ne ib 8 x 
Isachne confusa Ohwi 
Isachne firmula Buse 
Ischaemum arundinaceum F. Muell. ex Benth. 
Ischaemum australe R.Br. var. villosum Benth. 
Ischaemum decumbens Benth. ties de sat 
Ischaemum rugosum Salish. var. segetum (Trin.) Hack. : 
Tseilema dolichotrichum C. E. Hubbard .. ae oe x sits 
Tseilema fragile 8. T. Blake ae ot x es us x 


SKE GEO x X 


DS SOR OK OS OS OS 


Tseilema macratherum Domin fie a 
Tseilema membranaceum (Lindl.) Domin 
Tseilema vaginiflorum Domin : 
Iseilema windersw C. E. Hubbard 
Iseilema sp. 

Leptochloa brownti CO. E. Hubbard ; es Sy 42 
Leptochloa digitata (R.Br.) Domin Be ae iA x 
Lepturus repens (Forst. f.) R.Br. 

Lepturus xserophilus Domin 3 ate ar or 30 
tLolium rigidum Gaud. Ae se he ae Bae x 
Micraira subulifolia F. Muell. 

Neurachne alopecuroides R.Br. 

Neurachne mitchelliana Nees 

Neurachne muellert Hack. .. ke hide 
Neurachne munroi (F. Muell.) F. Muell. .. ote he vfs 
Ophiuros exaltatus (L.) Kuntze .. ie ie Br ee a x 
Oplismenus burmannit (Retz.) Beauv. We ge ae +e =e 
Oryza australiensis Domin 24 5a ty a ae wv: x 
Oryza rufipogon Griff. 

Panicum airoides R.Br. une Li sig i te BE ce 
Panicum australiense Domin a Ma as as x x x 
Panicum capillipes Benth... a Ss un We es a a 
Panicum cymbiforme Hughes 54 a aC ws x x x 
Panicum decompositum R.Br. be eae aN ue Xx x Xx 
Panicum delicatum Hughes oe oF at A: 

Panicum effusum R.Br... Be an x o 6 8 
Panicum majusculum F. Muell. ex ‘Benth. ae the Sad ws x 
Panicum pauciflorum R.Br. 

Panicum seminudum Domin ; he *iks oe we a se 
Panicum trachyrhachis Benth.  .. sie Hi a be ys Xx 
Panicum trichoides Swartz. . 

Panicum white J. M. Black 

Paractaenum novae-hollandiae Beauv. 
Paspalidium clementii (Domin) C. E. Hubbard - 
Paspalidium constrictum (Domin) C. KE. Hubbard ee = 9 afi eo 
Paspalidium distans (Trin.) Hughes A : i Se ss ot x 
Paspalidium gracile (R.Br.) Hughes ae Be i ote x te ois 
Paspalidium jubiflorum (Trin.) Hughes of a as ote x x 
Paspalidium rarum (R.Br.) Hughes as a: a x Xx x x 
Paspalidium retiglume (Domin) Hughes a ne af x oe 
Paspalum orbiculare Forst. f. i he af oF ds 2 i Xx 
Pennisetum arnhemicum F. Muell. ee i Be Bt obs x ate 
*Pennisetum polystachyon (Thell.) Schultz Se 48 ae er ft x 
Perotis rara R.Br. .. ae ate a Bi aft x x x x 


OOS OK x XX 

x x 
Ss OK NEES 
aD nie ee ON NEON 


SOK OS OK ORG OS OK EGS GOS OES OS OS 


xXXKX! 


PROCEEDINGS OF THE LINNEAN Society or NEw SourH Wates. Vou. 96, Part 4 


G. M. CHIPPENDALE 223 


C.A. Ba VER AD Darcie 


Pheidochloa gracilis S.T. Blake . ae a Te 
Phragmites karka (Retz.) Trin. ex Steud. i t33 x 
Plagiosetum refractum ee Muell.) Benth. a she x re ate 
Plectrachne bynoet C. EK. Hubbard oe he ae Ne: i x 
Plectrachne melvillei C. E. Hubbard Ae Pa Me x te 
Plectrachne pungens (R.Br.) C. E. Hubbard var. pungens x x x x 
Plectrachne pungens (R.Br.) C. E. Hubbard var. callosum 
C. E. Hubbard .. a ae ne ae a a Me: x 
Plectrachne schinzii Henr. .. ae ae ee Re x x x 
Pseudopogonatherum contortum (Brongn.) A. Camus 
Pseudopogonatherum irritans (R.Br.) A. Camus. . ae ae 
Pseudoraphis spinescens (R.Br.) J. Vickery ie iA x 
*Rhynchelytrum repens (Willd.) C. E. Hubbard .. a: x a ae 
Rottboellia formosa R.Br. .. ae se Sts Ea ae ee x 
Sacciolepis indica (l.) Chase ats 
Sacciolepis myosuroides (R.Br.) A. Camus ar eve i 
Schizachyrium fragile (R.Br.) A. Camus. as x x x 
Schizachyrium pseudeulalia (Hosokawa) S. T. Blake 
Sclerandrium grandifiorum 8.T. Blake .. a= 
Sclerandrium trancatiglume (F. Muell. ex Benth) Stapf 
et C. E. Hubbard ae ae ad a 
Sehima nervosum (Rottl.) Stapf .. = ae E. Sr x 
Setaria adhaerens (Forsk.) Chiov. ae ae 
Setarta apiculata (Seribn. et Merr.) K. Schum. aie e ahs 
Setaria brown Herrm. .. ae ie a aie x x 
Setaria dielsia Herrm. Ze ae ae ie ae x 
Setaria glauca (L.) Beauv. 
Setaria oplismenoides Herrm. Se te 58 a 
Setaria viridis (L.) Beauv. é Si eis x an : 
Sorghum australiense Garber et Snyder a = oe! x x 
Sorghum intrans F. Muell. ex Benth. 
Sorghum laxiflorum F.M. Bail. 
Sorghum macrospermum Garber mae a oe — uid 
Sorghum matarankensis Garber et Snyder a Ate at a x 
Sorghum plumosum (R.Br.) P. Beauv. ... ore an x x x 
Sorghum stipoideum (wart et White) C. A. Gardn. et 
C. E. Hubbard .. oe ao ae a ue 
Sorghum sp. aff. angustum 
Sorghum sp. ae oe Ae oe 
Spathia neurosa Ewart et “Archer ay ahs ae 4 a? 
Spinifex longifolius R.Br. a wie ea a we ae SK 
Sporobolus actinocladus (F. Muell.) F. Muell. 
Sporobolus australasicus Domin oe 
Sporobolus carolt Mez ah es a oe = 
Sporobolus mitchellii ee) C. E. Hubbard ex S.T. 
Blake = Sie oe ak x 
Sporobolus pulchellus R.Br. 
Sporobolus virginicus (L. ie Kunth. as Be ae 
Sporobolus sp : : ae Re ahs x 
Stipa scabra taney Pe ae aS x 
Thaumastochloa major S. T. Blake ey a 
Thaumastochloa pubescens (Domin) C. EH. Hubbard 
Themeda arguens (L.) Hack. : ae es ee 3G 
Themeda australis (R.Br.) Stapf : Be x <i ae 
Themeda avenacea (F.Muell.) Maid. et Betche | is >< S< 4 
Thuarea involuta (Forst. f.) R. et S. ss 
Tragus australianus 8. T. Blake x 
Triodia basedowiw EK. Pritz. au a Ps aa x 
Triodia clelandii N.T. Burbidge eh ws ate X 
Triodia hubbardit N. T. Burbidge is ae x ee by 
Triodia intermedia Cheel .. : ao ee Ec a ie S< 
Triodia inutilis N.T. Burbidge .. ies Bt om He XS x 
Triodia trritans R.Br. var. irritans ise Sc : 
Triodia longiceps J. M. Black F 2 Ne Bid a 
Triodia microstachya R.Br. ae Ex he é% ue ee x Xx 
Triodia procera R.Br. So a ate Hs ie Lt a x x 


x 
x 


OM XOX 
xX X x xXXXKXK KM KX: 


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x X 


PROCEEDINGS OF THE LINNEAN Socrpety or New SoutH Wates. Vou. 96, Part 4 


224 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Triodia pungens R.Br. var. linearis N.T. ees oy oe x 
Triodia pungens R.Br. var. pungens .. a6 x Xx 
Triodia racemigera C. A. Gardn. am 

Triodia roscida N.T. Burbidge .. Me we a et 

Triodia spicata N.'T. Burbidge .. bi are a: x 

Triodia stenostachya Domin : 

Triodia wiseana C. A. Gardn. var. wiseana ffs 
Tripogon loliiformis (F. Muell.) C. E. Hubbard 
Triraphis mollis R.Br. a ae Au bc 
Uranthoecium truncatum (Maid. et Betche) Stapf 
Vetiveria elongata (R.Br.) Stapf ex C. HE. Hubbard 
Vetiveria paucifiora S.T. Blake .. ats wt 
Whiteochloa semitonsa (F. Muell. ex Benth. ) C. E. Hubbard 
Xerochloa barbata R.Br. ae: 3S: ae on Bs ve out 
Xerochloa imberbis R.Br. .. nia be ate ce a oe x 
Xerochloa laniflora Benth.. : ae oe x ah x 
Zygochloa paradoza (R.Br.) ‘8. T. Blake tte if x 


A SON DN eS ee SNC 


5 IS eS OS ON 
x 


HN DN ONES OSG OS OSG Xe 


CYPERACEAE 

Arthrostylis aphylla R.Br. ae see ae aa ok Ws ay 
Bulbostylis barbata (Rottb.) C. B. Clarke a ae, S< x x 
Bulbostylis turbinata 8. T. Blake 6 mies ae x 

Caustis flecuosa R.Br. BE 4 

Cladium glomeratum R.Br... 

Cladium procerum 8. T. Blake 

Crosslandia setifolia W. V. Fitzg. 

Cyperus albomarginatus Mart. et Schrad. ex Nees 

Cyperus angustatus R.Br. 

Cyperus aquatilis R.Br. 

Cyperus armstrongii Benth. 

Cyperus articulatus L. : o6 26 do oc ot br ae 
Cyperus bifaz C. B. Clarke a Hh: i “m x x Xx 
Cyperus bulbosus Vahl ie x 

Cyperus castaneus Willd. var. brevimucronatus Kukenth. 

Cyperus compressus L. : a0 5 i ae ba 

Cyperus conicus (R.Br. a Boeck. .. Bis ee af Xx 

Cyperus cristulatus 8.T. Blake .. $s a ba ae 

Cyperus cunninghami (C. B. ee C. A. Gardn. Ye x 

Cyperus cuspidatus H.B.K. : boy a ae a a as 
Cyperus dactylotes Benth. Sk xe oe vie x Xx Xx 
Cyperus decompositus (R.Br.) F. Muell. “a cee Ne a 
Cyperus difformis L. Re oe Ko at - x x 3 
Cyperus diffusus Vahl 

Cyperus diphyllus Retz. .. om rs bes Ae tits 

Cyperus fasciculigerus (F. Muell.) Domin Lhe o% x a 

Cyperus gilesiz Benth. ae ce a a bis be x 

Cyperus gunni Hook. us wa as a ae x aM a 
Cyperus haspan L. 5 ne oe iG Af i me x 
Cyperus holoschoenus R.Br. ah Bes an Sc a x x 
Cyperus iria L. a be Si oe Xx x 

Cyperus txiocarpus F. Muell. on ae ae os x : ee 
Cyperus microcephalus R.Br. oP See 4 Ae mae Xx x 
Cyperus pennatus Lam. .. 3 ae at we = a x 
Cyperus platystylis R.Br. .. 

Cyperus polystachyos Rottb. aud sdf ae vee th at 
Cyperus pulchellus R.Br. .. = Aus us a te: bs x 
Cyperus pygmaeus Rottb. .. fe, ms ve dc < £8 x 
Cyperus ramos Kukenth.. . 

*Cyperus rotundus lL. me oe oa os as 

Cyperus rutilans (C. B. Clarke) Maid. et Betche re x 

Cyperus scariosus R.Br. ae ae Si i a a As 
Cyperus sporobolus R.Br. .. 58 or os ae +e x 
Cyperus squarrosus L. Bi: a4 2h i he x 

Cyperus stoloniferus Retz. : ss Me 

Cyperus subpinnatus WKukenth. var. subpinnatus 4 

Cyperus subpinnatus Kukenth. var. subrigidellus Kukenth. x 


XG NI 


MK Ge KKK MKS ROKK SG Bt PS OS SOc. ON Osos ON ON ON 


xX 5 


Nts 


PROCEEDINGS OF THE LINNEAN SocretTy or NEw SoutH WALES, Vot. 96, Part 4 


G. M. CHIPPENDALE 


225 


C.A. 


13) 51 


V.R.D. D. & G. 


Cyperus tegetuformis Roxb. 
Cyperus tenuispica Steud. 

Cyperus trinervis R.Br. 

Cyperus tuberosus Rottb. 

Cyperus vaginatus R.Br. .. ses 
Cyperus victoriensis C. B. Clarke 
Cyperus xserophilus Domin. . 


Cyperus sp. 
Cyperus sp. 


Diplacrum pygmaeum (R. Br.) Nees ex Boeck. 
Eleocharis atropurpurea (Retz.) Kunth. .. 
Eleocharis brassi 8. T. Blake ; 
Hleocharis caribaea (Rottb.) 8. T. Blake ; 
Hleocharis dulcis (Burm. f.) Trin. ex Hensch. .. 
Eleocharis nigrescens (Nees) Steud. 

Eleocharis pallens S.T. Blake 

Eleocharis sphacelata R.Br. fie 

Eleocharis spiralis (Rottb.) R. et S. 


Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimobristylis 
Fimbristylis 
Fimbristylis 
Fumbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Pimbristylis 


PROCEEDINGS OF THE LINNEAN SocreTy oF New SoutH WaALEs, 


acicularis R.Br. 

acuminata Vahl 

allenttd Turrill 

argentea Vahl ; 
arthrostyloides W.V. Fitzg. . 
bisumbellata (Forsk.) Bubani_ 
caespitosa R.Br. ae 
cardiocarpa F. Muell. 
cephalophora F. Muell. 
compacta Turrill 

corynocarya F. Muell. 
cymosa R.Br. 

cyperoides R.Br. 

debilis F. Muell. 

densa 8S. T. Blake 

denudata R.Br. 

depauperata R.Br... 
diphylla (Retz.) Vahl 


dipsacea (Rottb.) Benth. ox C. B. Clarke 
ferruginea Vahl a: : 5 
furva R.Br. 


humilis 8. T. Blake 
leucocolea Benth. 
littoralis Gaud. 
macrantha Boeck. 
macrostachya Boeck. 
microcarya FE. Muell. 
miliacea Vahl 
monandra F. Muell 
multifolia Boeck. 

nuda Boeck. ue 
nutans (Retz.) Vahl 
obtusangula F. Muell. 
oligocephala W. V. Fitzg. 
oxystachya FE. Muell. . . 
pallida 8. T. Blake 
paucifiora R.Br. oe 
phaeoleuca S. T. Blake 
pterygosperma R.Br... 
punctata R.Br. 
quinquangularis Kunth. 
rara R.Br. 

recta KF. M. Bail. 
rhyticarya F.Muell. .. 
schoenotdes (Retz.) Vahl 
schulizit Boeck. : 
sericea R.Br. 

setacea Benth. 


5 oN OS ON ON ON G 


x XX; 


xX! 


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SPOS ROR 


Vou. 96, Part 4 


226 


NORTHERN 


TERRITORY PLANTS 


C.A. 


Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Fimbristylis 
Pimbristylis 
Fimbristylis 
Pimbristylis 
Fimbristylis s 


solidifolia F. Muell. 
sphaerocephala Benth. 

spiralis R.Br.. A 
SQUATTOSA Vahl. var. squarrosa 


squarrosa Vahl var. esquarrosa Makino .. x 


squarrulosa F. Muell. 
stellata S. T. Blake 
tetragona R.Br. 
trachycarya FF. Muell. 
trigastrocarya EF. Muell. 
ayridis R.Br. F 
sp. aff. dichotoma 


Fuirena ciliaris (L.) Roxb. 
Fuirena incrassata 8. T. Blake 
Fuirena umbellata Rottb. 


Lepironia articulata (Retz.) Domin 


Lipocarpha microcephala (R.Br.) Kunth. 


Rhynchospora 
Rhynchospora 
Rhynchospora 
Rhynchospora 
Rhynchospora 
Rhynchospora 


affinis W.V. Fitze. 
heterochaeta 8. T. Blake 
longisetus R.Br... 
oterochaeta F. Muell. 
rubra (Lour.) Makino 
tenutfolia Benth. 


Schoenus falcatus R.Br. 
Schoenus sparteus R.Br. 
Scirpus articulatus L. 
Scirpus debilis Pursh. ; : 
Scirpus dissachanthus 8. T. Blake 
Scirpus grossus L..f. 3h: 
Scirpus laevis 8. T. Blake .. 
Scirpus lateriflorus Gmel. 
Scirpus litoralis Schrad. 
Scirpus maritimus L. 

Scleria brownit Kunth. 

Scleria ciliaris Nees 

Scleria hebecarpa Nees 

Scleria lithosperma Willd. .. 
Scleria margaritifera Willd. 
Scleria novae-hollandiae Boeck. 
Scleria pygmaea R.Br. 

Scleria rugosa R.Br. 

Scleria tricuspidata 8S. T. Blake 
Tricostularia fimbristyloides (F 


PALMAE 
Carpentaria acuminata (Wendl. et Drude) Bece. 


Corypha elata Roxb. 

Hydriastele wendlandiana Wendl. et Drude 
Kentia ramsayi (Becc.) Becc. 

Tivistona benthamu ¥F.M. Bail. 

Tivistona humilis R.Br. 

Livistona loriphylla Bece. 

LTivistona mariae F. Muell. 

Nypa fruticans Wurmb. .. 

Ptychosperma elegans (R.Br.) Bk 


ARACEAE 


PROCEEDINGS OF THE LINNEAN Society or NEW 


Amorphophallus galbra F. M. Bail. 
Amorphophallus variabilis Bl. 

Pistia stratiotes L. .. 

Raphidophora australasica ¥. M. Bail. 
Typhonium angustilobum F. Muell. 
Typhonium browni Schott. 
Typhonium liliifolium Schott. 
Typhonium oblongifolium Engl. 


F. Muell.) Benth. 


5 wes ON ek G 


x 


SoutH WALES, 


Vou. 


96, 


5 os OS Oxo 


XxX: 


Part 4 


Sees 


xXxXXX: SRK ee Re OS XN RUS Rs ee OU ROG og 


KMXKKKKKXKXK: 


xX XX XK XK IS OR OG OSPR OS OOS ON 


iG 


G. M. CHIPPENDALE 


FLAGELLARIACEAE 


Flagellaria indica L. 


RESTIONACEAE 


Leptocarpus elatior R.Br. .. 
Leptocarpus spathaceus R.Br. 


CENTROLEPIDACEAE 


Centrolepis banksw (R.Br.) R. et 8. 
Centrolepis exserta (R.Br.) R. et 8. 
Centrolepis polygyna (R.Br.) Hieron 
Centrolepis pusilla (R.Br.) R. et S. 


XYRIDACEAE 


Xyris complanata R.Br. 
Ayris juncea R.Br. 
AXyris paludosa R.Br. 
Xyris pauciflora Willd. 


ERIOCAULACEAE 


Hriocaulon australe R.Br. 

Eriocaulon bifistulosum van Heurck et J. Muell. 

Hriocaulon cinereum R.Br. : % 

EHriocaulon graphitinum F. Muell. et Tate ex Ewart et 
Cookson 2 

Eriocaulon heterogy, ynum F. Muell. 

Hriocaulon scartosum R.Br. 

Eriocaulon schultzii Benth. 

Eriocaulon tortuosum FE. Muell. 

Eriocaulon sp. 


COMMELINACEAE 


Cartonema parviflorum Hassk. 
Cartonema spicatum R.Br. 

Commelina cyanea R.Br. 

Commelina ensifolia R.Br. 

Commelina lanceolata R.Br. 
Commelina undulata R.Br. 

Cyanotis axillaris (L.) D.Don .. 
Murdannia gigantea (R.Br.) Bruckn. 
Murdannia graminea (R.Br.) Bruckn. 
Murdannia vaginata (L.) R.Br. 


PONTEDERIACEAE 


Monochoria cyanea F. Muell. 


PHILYDRACEAE 


Philydrum lanuginosum Banks et Soland. ex Gaertn. 


JUNCACEAE 


Juncus sp. 
Juncus sp. 


RoOxXBURGHIACEAE 


Roxburghia javanica Kunth. 


LILIACEAE 


Anguillaria dioica R.Br. . 

Asparagus racemosus Willd. 

Caesia setifera Baker a 

Corynotheca lateriflora (R. Br.) Benth. 

Dianella caerulea Sims 56 

Dianella ensifolia Red. 

Dracaena angustifolia Roxb. ue o¢ st a 
Lomandra leucocephala (R.Br.) Ewart ssp. robusta A. Lee 
Lomandra multiflora (R.Br.) J. Britt. 8 ae 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH WaAtLEgES, VOL. 


Ns V.R.D. D. & G. 


96, 


mS OS OS ON G OS ON oleN eS 26 ON OSes ON 0 xX X X xX XK XX 


x 


RN KOK OK KE 


Oa: 


Part 4 


NORTHERN TERRITORY PLANTS 


C.A. 

Lomandra patens A. Lee .. Ae ee te 4 x 
Smilax australis R.Br. 

Smilax latifolia R.Br. 

Sowerbaea alliacea F. Muell. 

Thysanotus chinensis Benth. A 
Thysanotus exiliflorus F. Muell. x 
Thysanotus tuberosus R.Br. x 
Tricoryne elatior R.Br. x 
Xanthorrhoea thornton Tate x 


B.T. 


V.R.D. D.& G. 


HAEMODORACEAE 


Haemodorum brevicaule F. Muell. 
Haemodorum coccineum R.Br. ; 
Haemodorum ensifolium F. Muell. 
Haemodorum leptostachyum Benth. 
Haemodorum parviflorum Benth. .. 
Haemodorum subvirens F. Muell. 


AMARYLLIDACEAE 


Crinum asiaticum L. var. asiaticum : 

Crinum asiaticum L. var. angustifolium Benth. ae es 
Crinum flaccidum Herbert ae a ae or x 
Crinum uniflorum F. Muell. 

Curculigo ensifolia R.Br. 


TACCACEAE 


Tacca leontapetaloides (L.) Kuntze 


DIOSCOREACEAE 


Dioscorea bulbifera L. 5 

Dioscorea sativa L. var. sativa ; oe 
Dioscorea sativa L. var. rotunda F.M. Bail. 
Dioscorea transversa R.Br. 


ZINGIBERACEAE 


Curcuma australasica Hook. f. 


BuRMANNIACEAE 


Burmannia juncea Sol. 


ORCHIDACEAE 


Cymbidium canaliculatum R.Br. ar. barrettii W. H. 
Nicholls Be ad 
Dendrobium dicuphum F. Muell. 
Dendrobium johannis Reichb. f. 
Dipodium stenocheilum O. Schwarz 
Eulophia holtzei F. Muell. . . 
Hulophia venosa Reichb. f. 
Geodorum pictum (R.Br.) Lindl. 
Habenaria elongata R.Br. at 
Habenaria ferdinandi Schlechter . . 
Habenaria holtzer F. Muell. Fe 
Habenaria hymenophylla Schlechter 
Habenaria ochroleuca R.Br. 
Luisia teretifoia Gaud. 
Nervilia sp. 
Spiculaea sp. 


CASUARINACEAE 


Casuarina cunninghamiana Miq. om ae Pr 
Casuarina decaisneana F. Muell. aft a O10 x 
Casuarina equisetifolia Forst. et Forst. f. 


PIPERACEAB 


Peperomia pellucida (L.) H.B.K... 
Piper holtzec F. Muell. x 
Piper novae-hollandiae Miq. 


x 
x 
x 
5 x 
oti x 
x x 
x 
x 
3: x 
x 
x x 
x 
x 
x 
x x 
x 
Si 
x 
x 
x 
x 
x 
x 
Se 
x 
x 
x 
x 
x 
x 
xe 
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x 
x 
x x 
x 
x 
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PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 229 


C.A. lef ho \WolsD)S 1D), a5 (Gr 


ULMACEAE 
Celtis philippensis Blanco ie re ae it she a0 x x 
Trema amboinensis (Willd.) Bl. .. a a ae Ai 46 Bc x 
Trema aspera (Brongn.) BI. Be 2/5 ie ie x x Xx 


MORACEAE 

Antiaris toxicaria Lesch. var. macrophylla (R.Br.) Corner... hs Se x 
Fatoua pilosa Gaud. te Pee a ais ae are ws x 
Ficus benjamina L. var. benjamina = a x. on 5h x 
Ficus coronulata F. Muell. fa ai se os ae ne x 

Ficus hispida L.f. var. hispida ae “his a a a x 
Ficus lacor Buch. Ham. .. a “8 ve a es “i x 
Ficus leucotricha Miq. var. leucotricha a 6 x 
Ficus leucotricha Miq. var. megacarpa F. Muell. ex - Corner ae x v 

Ficus obliqua Forst. f. var. obliqua : ae 50 a Xx 
Ficus opposita bees var. indecora (A. Cunn. ex Mia.) 

Corner .. 3 i: St se x 
Ficus opposita Mig. var. micracantha (Miq.) Corner .. oD ot x x 
Ficus platypoda (Miq.) A. Cunn. ex pet var. angustata 

(Miq.) Corner .. ore Pe as x 
Ficus plat uo pede (Miq.) A. ‘Cunn. ex Mia. var. cordata 

Specht ee oe x 
Ficus Pode (Miq. A. Cunn. ex - Miq. var. minor Benth. x 3 ¢ x 
Ficus platypoda (Miq.) A. Cunn. ex pee var. a eo ee x we 
Ficus racemosa L. var. racemosa ae wt x 
Ficus scobina Benth. é He ye 
Ficus superba Miq. var. henneana (Miq.) Corner 
Ficus virens Ait. 

Malaisia scandens (Lour.) ‘Planch. 


MK OK OK RK a 


URTICACEAE 
Parietaria debilis Forst. f. oe af ae ag x 
TUrtica urens L. of B09 a ak 56 o 0 x 


PROTEACEAE 
Banksia dentata L.f. 
Grevillea agrifolia A. Cunn. ex R. Br. 
Grevillea angulata R.Br. .. 
Grevillea dimidiata F. Muell. isd wit a4 ss bis 
Grevillea dryandrt R.Br... Bm ae ae oe A x 
Grevillea eriostachya Lindl. ae se ae He < 6 
Grevillea goodw R.Br. A me os & vA $2 ti ee x 
Grevillea heliosperma R.Br. au ae ae of oe x 
Grevillea juncifolia Hook. ae ae x x 
Grevillea leucadendron A. Cunn. ex oe Br. 
Grevillea mimosoides R.Br. ns Ses exe ae 
Grevillea nematophylla F.Muell. .. ag ae 2 x 
Grevillea pachypoda O. Schwarz <a a a ni 0 
Grevillea parallela Knight . . ae ag a ae ks x 
Grevillea pteridiifolia Knight 
Grevillea pungens R.Br. 
Grevillea refracta R.Br. 
Grevillea stenobotrya KF. Muell. 
Grevillea striata R.Br. 
Grevillea wickhamw Meissn. 
Hakea arborescens R.Br. .. 
Hakea chordophylla F. Muell. 
Hakea cunninghamiu R.Br. 
Hakea divaricata L. Johnson 
Hakea leucoptera R.Br. 
Hakea macrocarpa A. Cunn. ex R.Br. 
Hakea morrisoniana W.V. Fitzg.. 
Hakea multilineata Meissn. var. grammatophylla B. Muell. 
Hakea suberea S. Moore ae oe : 
Hakea sp. 
Helicea australasica ‘F. Muell. : ae af ee fe 
Persoonia falcata R.Br... orc ue 5A ee ee 0 “< 
Stenocarpus cunningham R.Br. .. 


SOK 6 So 6 KAR KM 
5 OS OX OX 


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PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WaAtss, Vou. 96, Part 4 


230 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. &G. 


LORANTHACEAE 

Amyema benthamw (Blakely) Dans. ; = au oe bi oe; 
Amyema bifurcatum (Benth.) Tiegh. se Bie fs x x x 
Amyema brevicaule Barlow 
Amyema conspicuum (FE. M. Bail. ) Dans. ssp. obscurinerve 

Barlow .. fee ae Be aa ar ah ee x 
Amyema fulviflorum Barlow ; oh a Spe sci x 
Amyema gibberulum (Tate) Dans. var. gibber elum Se x ae ot Be 
Amyema haematodes (O. Schwarz) Dans. oy ae ES af oe x 
Amyema herbertianum Barlow .. ae We ie £4 i bgt x 
Amyema hillianum (Blakely) Dans. = Be He x 
Amyema mackayense (Blakely) Dans. ssp. cycnei-sinus 

(Blakely) Barlow 3 ae a axe as sa ey ys « 
Amyema maidenir (Blakely) Barlow ssp. maiden... x ~ Ae ae 
Amyema miquelit (Lehm. ex Mia.) Tiegh. ne ihe x SP se x 
Amyema miraculosum One) Tiegh. Ssp. boormania 

(Blakely) Barlow : or oe st x pe ae Se 
Amyema pendulum (Sieb. ex Spreng. ) Wigan... ot te 5, x x 
Amyema preissii (Miq.) Tiegh. .. ie x a Bac 
Amyema quandang (Lindl.) Tiegh. var. quandang ae x ey: x 
Amyema sanguineum (F. Muell.) Dans. var. pulcher 

(Ewart) Barlow ie gts ve ae ais: re a me: x 
Amyema sanguineum (F. Muell.) Dans. var. sanguineum x of x 
Amyema thalassitum Barlow iS he at me 4 
Amyema villiflorum (Domin) Barlow ssp. tomentellum 


x XX 


(Blakely) Barlow Ae tt ope x 
Amyema villiflorwm (Domin) Barlow ssp. villiflorum x 
Decaisnina brittenii (Blakely) Barlow a x 
Decaisnina petiolata (Barlow) Barlow ae Be x 
Decaisnina signata (F. Muell. ex Benth.) Barlow ssp. 

cardiophylla (Domin) Barlow NS = x 
Decaisnina signata (F. Muell. ex Benth. .) Barlow ssp. 

signata .. oc ae a2 re 3 20 26 x 
Dendrophthoe acacioides (A. Cunn. ex Benth.) Tiegh. 

var. acacioides .. ie es zs es 50 x x 
Dendrophthoe acacioides (A. Cunn. ex Benth.) Tiegh. 

var. longifolia (Specht) Barlow uM he a fe sks x 4 
Dendrophthoe falcata (L.f.) Ettingsh. .. a: Bo ae 55 x x 
Dendrophthoe glabrescens (Blakely) Barlow re oe x 
Dendrophthoe odontocalyx (F. Muell. ex ge Tiegh. x 
Diplatia furcata Barlow .. : Fe He a x 
Diplatia grandibractea (F. Muell.). Tiegh. : x x x 
Lysiana exocarpt (Behr) Tiegh. ssp. diamantinensis 

(J. M. Black) Barlow .. fe ae x 
Lysiana murrayt (F. Muell. et Tate) Tiegh. Se x 
Lysiana spathulata (Blakely) Barlow ae! parvifolia 

Barlow .. fe x =: Ae 
Lysiana spathulata (Blakely) Barlow ssp. spathulata 6 >< x x x 
Lysiana subfalcata (Hook.) Barlow eee, ae Ses x re x an 
Viscum flezicaule Barlow .. an Pe Bs a x 

SANTALACEAE 
Anthobolus leptomerioides F. Muell. ae ae x he ae a 
Anthobolus filifolius R.Br. Be a we sa Be De on x 
Hzxocarpos latifolius R.Br. es she oe ay as BS x x 
Haocarpos sparteus R.Br. .. x ne : 
Santalum acuminatum (R.Br.) A. DOs x 
Santalum lanceolatum R.Br. var. angustifolium Benth. x oe 
Santalum lanceolatum R.Br. var. lanceolatum ae x x x x 
Santalum ovatum R.Br... ua ee 36 eae Ae wes se x 
OPILIACEAE 
Cansjera leptostachya Benth. a on ae ay hy Le 2% x 
Opilia amentacea Roxb. .. oe ans ts La nie on x Xx 
OLACACEAE 
Olax aphylla R.Br. ane rs a Se te a aif ae x 


PROCEEDINGS OF THE LINNEAN Society or New SourH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 231 


C.A. BT. V.R. D. D. & G. 


ARISTOLOCHIACEAE 
Aristolochia thozetis F. Muell. 


POLYGONACEAE 
+Emex australis Steinh. oe as De bu x 
Muehlenbeckia cunninghamii (Meissn.) F. Muell. x » x A 
Polygonum attenuatum R.Br. Ae te an x x x 
+Polygonum aviculare L.  . x 
Polygonum lapathifolium Tbe, x 


Polygonum orientale L. he si 1h oe ae ne Re eS x 
Polygonum plebecum R.Br. 
Polygonum serrulatum Lag. 
Rumex crystallinus Lange . . 
*Rumex vesicarius Li. 


XX XK X 


CHENOPODIACEAE 

Arthrocnemum halocnemoides Nees var. halocnemoides 

Arthrocnemum halocnemoides Nees var. pergranulatum 
J. M. Black ae 34 iG ips 5 Me ae se x 

Arthrocnemum leiostachyum (Benth.) Paulsen 

Atriplex angulata Benth. .. 

Atriplea conduplicata F. Muell. 

Atriplex elachophylla F. Muell. 

Atriplex holocarpa F. Muell. 

Atriplex incrassata F. Muell. 

Atriplec limbata Benth. 

Atriplex lindleyi Moq. 

Atriplex muellert Benth. 

Atriplezr nummularia Lindl). bi 

Atriplex pseudocampanulata Aellen 

Atriplex semibaccata R.Br. 

Atriplex spongiosa F. Muell. ; 53 

Atriplex turbinata (R. H. Anderson) Aellen 

Atriplex velutinella F. Muell. ee 

Atriplex vesicaria Heward ex Benth. 

Babbagia acroptera F. Muell. et Tate 

Babbagia dipterocarpa F. Muell. 

Bassia anderson Ising 4 

Bassia bicornis (Lindl.) F. Muell. var. bicornis 

Bassia bicornis (Lind].) F. Muell. var. horrida (Domin) 
C. T. White ay Be 

Bassia biflora (R.Br.) F. Muell. var. biflora 

Bassia birchit (F. Muell.) F. Muell. : 

Bassia calcarata Ising is ae 

Bassia chippendalei Ising 

Bassia clelandw Ising bef an 

Bassia convexula R.H. Anderson. . 

Bassia cornishiana F. Muell. 

Bassia costata R. H. Anderson 

Bassia decurrens J. M. Black 

Bassia diacantha (Nees) F. Muell. 

Bassia divaricata (R.Br.) F. Muell. 

Bassia eremaea Ising ee a: 

Bassia eriacantha (F. Muell.) R. H. Anderson 

Bassia glabra (F. Muell.) F. Muell. 

Bassia intricata R. H. Anderson .. 

Bassia johnsoni Ising : oe 

Bassia lanicuspis (F. Muell.) F. Muell. 

Bassia limbata J. M. Black : 

Bassia longicuspis F. Muell. 

Bassia luehmannii F. Muell. 

Bassia obconica Ising 

Bassia obliquicuspis R. H. ‘Anderson 

Bassia paradoxa (R.Br.) F. Muell. 

Bassia parallelicuspis R.H. Anderson 

Bassia patenticuspis R. H. Anderson 


Z 
x 


XKXKXKKKKKKKKKK KKK KKK 


He KG OK OK OK OS OK OK OK ON OK OK OK OK OK OS OR OS OK ERK OS 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Wass, Vou. 96, Part 4 


232 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Bassia a eel es (F. Muell.) F. Muell. var. lanata 
Ising ‘ ws he ee a ahs 

Bassia quinquecuspis (F. Muell.) F. Muell. var. villosa 
(Benth.) J. M. Black .. ae 

Bassia sclerolaenoides (F. Muell.) F. Muell. 

Bassia urceolata \sing 

Chenopodium anidioph yllum ‘Aellen 

Chenopodium auricomum Lindl. .. ae 

Chenopodium cristatum (F.Muell.) F.Muell. .. 

Chenopodium desertorum (J.M. Black) J.M. Black  .. 

Chenopodium melanocarpum (J.M. Black) J. M. Black 

tChenopodium murale L.  . 

Chenopodium nitrariaceum (F. Muell. ) F. Muell. ex “Benth. 

Chenopodium pumilio R.Br. : 

Chenopodium rhadinostachyum F. Muell. 

Dysphania littoralis R.Br. 

Dysphania plantaginella F. Muell. 

Dysphania simulans F. Muell. et Tate ex : Tate 

Enchylaena tomentosa R.Br. se ae 

Kochia aphylla R.Br. 

Kochia appressa Benth. .. 

Kochia astrotricha L. Johnson 

Kochia brevifoia R.Br... 

Kochia coronata J. M. Black 

Kochia enchylaenoides (J. M. Black) J. M. Black 

Kochia georger Diels Bic 

Kochia lanosa Lindl. 

Kochia lobiflora F. Muell. 

Kochia planifolia F. Muell. 

Kochia scleroptera J. M. Black 

Kochia spongiocarpa F. Muell. 

Kochia tomentosa F. Muell. 

Kochia triptera Benth. 

Kochia villosa Lindl. 

Kochia sp. 

Kochia sp. . a6 or 56 

Pachycornia tenuis (Benth.) J. M. Black 

Pachycornia triandra (KF. Muell.) J. M. Black 

Rhagodia nutans R.Br. se Be 

Rhagodia parabolica R.Br.. 

Rhagodia spinescens R.Br... 

Salsola kali L. var. kala 

Salsola kali L. var. strobilifera Benth. 

Tecticornia cinerea (F. Muell.) Hook. f. et Jackson 

Threlkeldia inchoata (J.M. ee J.M. Black . 

Threlkeldia procerifiora F. Muell. z: 


x 


SS NE SK SS DK EK OR NE OR KR ORK KOK KK KKK KKK KKK KX BK KK KKK KK KK KX 
x 
x 
5 ek OS OS 6 


AMARANTHACEAE 

Achyranthes aspera L. 
Alternanthera anqustifolia R.Br. 
Alternanthera denticulata R.Br. .. Se 
Alternanthera micrantha (Benth.) Domin 
Alternanthera nana R.Br. evs shi site 3 x x 
Alternanthera nodiflora R.Br. ate 5 Bed 5.6 x x x 

*Alternanthera pungens H.B.K. : se Be x x 
Amaranthus cochleitepalus Domin ae aye a a x 
Amaranthus grandifiorus (J.M. Black) J.M. Black .. x Re at 
Amaranthus interruptus R.Br... ae 5 it Xx is x 
Amaranthus leptostachyus Benth. oy a at ts if, 
Amaranthus mitchelliia Benth. sa an ste a x x ae 
Amaranthus pallidifiorus F. Muell. O0 56 20 5c Ae x 

tAmaranthus viridis L. Ne ih i ae =< 

*Celosia argentea Ih. ue ay cd a he. ans a 
Gomphrena affinis F. Muell. ts ae ae a iY: ve x 
Gomphrena alba Farmar .. Ne ae aa a ok A a: 
Gomphrena brachystylis F. Muell. at ot ae Be, x x 


xX X X 

2X 

xX X X 
xX XK XK XK 


KX) 


ee 


XXxXXX: 


PROCEEDINGS OF THE LINNEAN SociETY or NEw SoutH WALES, Vot. 96, Part 4 


G. M. CHIPPENDALE 


C.A. Isto (VIR ID IDS 64 Gr 


Gomphrena canescens (Poir.) R.Br. 

Gomphrena conferta Benth. 

Gomphrena conica Spreng. 20 He 

Gomphrena cunninghamw (Moq.) Druce .. 

Gomphrena flaccida R.Br. 

Gomphrena humilis R.Br. 

Gomphrena lanata R.Br. .. 

Gomphrena parviflora Benth. ae 

Ptilotus aervoides (F. Muell.) F. Muell. 

Ptilotus aristatus Benl 2a 

Ptilotus arthrolasius F. Muell. A 1x ne 

Piilotus atriplicifolius (A.Cunn. ex Mog.) Benl var. 
atriplicifolius oe S¢ aye ig an: 

Ptilotus atriplicifolius (A. Cunn. ex Mog.) Benl. var. 
eldert (Farmar) Benl. as a ste it 

Ptilotus calostachyus (F. Muell.) F. Muell. var. calostachyus 

Piilotus calostachyus (F. Muell.) F.Muell. var. procerus 
(Diels) Benl By ae 

Ptilotus clementi (Farmar) Benl . 

Piilotus conicus R.Br. 

Ptilotus corymbosus R.Br. var. acutiflorus Benth. 

Ptilotus corymbosus R.Br. var. corymbosus 

Ptilotus decipiens (Benth.) C. A. Gardn. 

Piilotus dissitiflorus F. Muell. i 

Ptilotus distans (R.Br.) Poir. 

Ptilotus exaltatus Nees ex Lehm. : 

Pitlotus fusiformis (R.Br.) Poir. var. fusiformis an 

Ptilotus fusiformis (R.Br.) Poir. var. gracilis (R.Br.) Benl 

Ptilotus gaudichaudiu (Steud.) J.M. Black var. ee 
chaudiw .. te : 

Ptilotus helipteroides (F. Muell.) F. “Muell. oe 

Ptilotus incanus (R.Br.) Poir. var. elongatus Benl 

Ptilotus incanus (R.Br.) Poir. var. incanus 

Pilotus latifolus R.Br. var. latifolius ae 

Ptilotus latifolius R.Br. var. mator (C. A. Garda.) Benl 

Ptilotis leucocoma (Moq.) F. Muell. a 

Ptilotus lophotrichus Benl 

Prtilotus macrocephalus (R.Br.) Poir. 

Ptilotus murrayi F. Muell. : = 

Ptilotus nobilis (Lindl.) F. Muell. var. nobilis be 

Ptilotus obovatus (Gaud.) F. Muell. var. griseus Benl | 

Pitilotus obovatus (Gaud.) F. Muell. var. obovatus 

Ptilotus obovatus (Gaud.) F.Muell. var. parviflorus 
(Lindl.) Benl .. ie 5% oe a 

Ptilotus parvifolius (F. Muell.) F. Muell. om $e 

Piilotus polystachyus (Gaud.) F.Muell. emend Benl 
forma polystachyus  .. ar 50 ae at 

Ptilotus poalystachyus (Gaud.) F.Muell. emend Benl 
forma rubriflorus (Black) Benl a3 ee 

Pitilotus schwartz F.Muell. ex Tate a A 

Ptilotus spicatus F.Muell. ex Benth. ssp. leianthus 
(Benth.) Benl var. letanthus ae as 

Pitilotus spicatus F.Muell. ex Benth. ‘ssp. leianthus 
(Benth.) Benl var. longiceps Benl .. an is 

Ptilotus spicatus F. Muell. ex Benth. oe, spicatus 

Pupalia lappacea (l.) Juss. : ae 


NYCTAGINACEAE 


Boerhavia diffusa L. Shs 
Boerhavia repanda Willd. 
Pisonia aculeata L. 


GYROSTEMONACEAE 


Codonocarpus cotinifolius (Desf.) F. Muell. 
Gyrostemon australasicus (Moq.) Heimerl. 
Gyrostemon ramulosus Desf. 4 


PROCEEDINGS OF THE LINNEAN SocreTy or NEW 


x 
y. 
x x x 
x 
x es 
x x 
x 
x & 
x x x 
x 
x x 
x nt 
x 
x x x 
x x x 
x x 
x 
x a: 
ys x 
x 
x 
x 
x 
x x x 
x 
x x 
x ei 
x x 
M8 x 
x 
x x 
x x 
x 
x 
x x x 
x x x 
x 
x 
x 
x 


SoutH WALEs, VOL. 


S§ oMiek a 


xX XXX: 


CNOA eS 


96, Part 4 


NORTHERN 


TERRITORY PLANTS 


V.R.D. D. 


C.A. Bey & G. 
Gyrostemon sp. x 56 
Gyrostemon sp. x x 
SPHENOCLEACEAE 
Sphenoclea zeylanica Gaertn. 4 
AIZOACEAE 
Aizoon quadrifidum (F. Muell.) F. Muell. S< EY: 
Aizoon zygophylloides (F. Muell.) F. Muell. x M3 x 
Glinus lotoides L. .. ne ue ee x x x 
Glinus oppositifolius (L.) A.DC. .. x x x 
ilinus orygioides F. Muell. Xx A 
Macarthuria apetala Harv. ie x 
Mollugo cerviana (L.) Ser. ate >< oe : 
Mollugo molluginea (F. Muell.) Druce x x x Me 
Mollugo pentaphylla L. 5 x 
Sesuvium portulacastrum L. by x 
Trianthema oxycalyptra F. Muell. x af : a 
Trianthema pilosa F. Muell. x x x x 
Trianthema portulacastrum 1. x 
Trianthema triquetra Willd. var. " clavata (J. M. : Black) 
Eichler .. x x 
Trianthema tr iquetra Willd. var. triquetra x te x 
Zaleya galericulata (Melville) Eichler x x 
PORTULACACEAE 
Calandrinia balonensis Lindl. x 2 
Calandrinia eremaea Ewart x Ae 
Calandrinia gracilis Benth. Dy x 
Calandrinia polyandra Benth. x 
Calandrinia ptychosperma F. Muell. x 
Calandrinia pumila (F. Muell. ex Benth.) F. Muell. x a A. 
Calandrinia quadrivalvis KF. Muell. : a x x 
Calandrinia remota J. M. Black x 
Calandrinia spergularina F. Muell. x 
Calandrinia stagnensis J. M. Black x cc oe 
Calandrinia uniflora F. Muell. x x 
Portulaca bicolor F. Muell. use x x 
Portulaca digyna F. Muell. os x x 4% 
Portulaca filifolia F. Muell. x x x x 
Portulaca intraterranea J.M. Black x at ise ae 
Portulacu oleracea L. x x x x 
Portulaca oligosperma F. Muell. aR x x i. 
Portulaca CERES F. Muell. ex Benth. ae é x x 
Portulaca sp. : x E 
CARYOPHYLLACEAE 
+Cerastium aff. glomeratum Thuill . . x oe - 
Polycarpaea breviflora F. Muell. x x x 
Polycarpaea burtoniit F.M. Bail. oa ae ah Se 
Polycarpaea corymbosa (L.) Lam. x x * x 
Polycarpaea holtzii Maid. et Betche ae 5 x 
Polycarpaea involucrata F. Muell.. 2 P x iz x x 
Polycarpaea longiflora F. Muell. var. longiflsra 24 x x x 
Polycarpaea longiflora F. Muell. var. lewcantha Benth. x x 
Polycarpaea staminodina F. Muell. : . a x x 
Polycarpaea synandra ¥F. Muell. x x x 
Polycarpaea triloba Ewart et Cookson x - 
Polycarpaea violacea (Mart.) Benth. a x 
Polycarpon Te (L.) L. x 
TSpergularia sp. : My x 
NYMPHAEACEAE 
Nelumbo nucifera Gaertn. .. Se se . x 
Nymphaea gigantea Hook. var. gigantea ae a x x 
Nymphaea violacea Lehm. x x x 
PROCEEDINGS OF THE LINNEAN SocreTy or New Sours Wates, Vou. 96, Part 4 


G. M. CHIPPENDALE 


CERATOPHYLLACEAE 
Ceratophyllum demersum L. 


RANUNCULACEAE 
tAdonis aestivalis L. 


MENISPERMACEAE 
Hypserpa decumbens (Benth.) Diels : 
Pachygone ovata Miers ex Hook. f. et Thomas 


Stephania japonica ge Miers var. timoriensis 


(DC.) Forman 
Tinospora smilacina Benth. 


ANNONACEAE 
*Annona reticulata L. 
Polyalthia holtzeana F. Muell. gas 
Polyalthia nitidissima (Dene.) Benth. 
Popowia australis Benth. .. : 
Uvaria holtzet F. Muell. 
Uvaria membranacea Benth. 


MYRISTICACEAE 
Myrtistica insipida R.Br. .. 
Horsfieldia australiana 8. T. Blake 


LAURACEAE 
Cassytha filuformis L. 
Cassytha glabella R.Br. 
Cryptocarya cunningham Meissn. 
Intsea chinensis Lam. i 
Iitsea glutinosa (Lour.) C. B. Robinson .. 


HERNANDIACEAE 
Gyrocarpus americanus Jacq. 


PAPAVERACEAE 
+Glaucitum corniculatum (L.) Rudolph 
tPapaver hybridum L. SA ae 


CRUCIFERAE 

Arabidella trisecta (F.Muell.) Schulz 

Blennodia canescens R.Br. Fi ae 
Blennodia pterosperma (J. M. Black) J. M. Black 
*Brassica tournefortis Gouan. : a Ae 
+Capsella bursa-pastoris (L.) Medik. 
+Carrichtera annua (L.) Prantl —- a: 
Harmsiodoxa blennodioides (F. Muell.) Schulz 
Lepidium muelleri-ferdinandi Thell. : 
Lepidium oxytrichum Sprague 
Lepidium papillosum F. Muell. 
Lepidium rotundum (Desv.) DC. 
Menkea sphaerocarpa F. Muell. uf a 
Menkea villosula (F. Muell. et Tate) J. M. Black 
Phlegmatospermum cochlearinum (F. Muell.) Schulz 
+Sisymbrium erysimoides Desf. Se ai 
+Sisymbrium wvrio L. 
+Sisymbrium orientale L.  . 
Stenopetalum lineare R.Br. ex DC. 
Stenopetalum nutans F. Muell. 
Stenopetalum velutinum F. Muell. 


CAPPARIDACEAE 
Capparis lasiantha R.Br. ex DC. 
Capparis loranthifolia Lindl. var. loranthifolia 
Capparis mitchellii (Lindl. ex F. ae Benth. 
Capparis quiniflora DC. a; 


PROCEEDINGS OF THE LINNEAN SocretTy or New Sours Wa.eEs. VoL. 96, 


xX KKK KKK KKK KKK KKK XK XK XK XK Xe BK 


x XX 


235 


VRD. D. & G. 


x xX 


x KK KX 


x 


Part 4 


236 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Capparis re L. ssp. wumbellata (R.Br. ex DC.) 

Jacobs a 
Capparis spinosa L. var. nummularia De.) F. M. Bail. x Ae 
Capparis umbonata Lindl. x x 
Cleome ouwalidea F. Muell. 

Cleome tetrandra Banks ex DC. ae ie ae Gd és 
Cleome viscosa L. .. ee ies ie ae ah x < 
Justago cleomoides (F. Muell.) Kuntze 


5 OS 
x 


xxx xXx 
EXE KORE 


DROSERACEAE 
Drosera burmanni Vahl .. oy a, a ee x i xx 
Drosera indica L. .. Me sh me ats ae x x x 
Drosera petiolaris R.Br. ex DC. ie We bas it ee x 


xX XX 


BYBLIDACEAE 
Byblis liniflora Salisb. < of te ne aa ar he x % 


CRASSULACEAE 
Crassula colorata (Nees) Ostenf. .. a: ae a < 
Crassula purpurata (Hook. f.) Domin .. bye si x 


PITTOSPORACEAE 
Pittosporum ferrugineum Ait. we a oc a3 ao ovo xX 
Pittosporum melanospermum F. Muell. .. ‘its be bt WY x x 
Pittosporum phylliraeoides DC... ae a ae x aa < 


RosACEAE 
Parinari corymbosum (Bl.) Miq. .. ae a Ne Le sc 50 x 
Rubus hill F. Muell. ee aw ik oe of an af a) x 
Stylobasium spathulatum Desf. 


x 


MIMOSACEAE 

Acacia acradenia F. Muell. ae ay: or ¥e x Xx 
Acacia adsurgens Maid. et Blakely ae ae cs 4 4 x . 
Acacia alleniana Maid. .. ah a 3 of ea ah x 
Acacia amentifera F. Muell. whe a sa sat 6 x 
Acacia ancistrocarpa Maid. et Blakely ashe ae Ny < Xx xX 
Acacia aneura F. Muell. ex Benth. var. aneura 4 x 
Acacia aneura F. Muell. ex Benth. var. ee J. M. 

Black... : =; x 
Acacia argyraea Tindale 
Acacia arida Benth. : a be? a, st ic es Pe 
Acacia asperulacea F. Muell. ws a Bi x 
Acacia aulacocarpa A. Cunn. ex Benth. var. aulacocarpa of x x 
Acacia aulacocarpa A. Cunn. ex Benth. var. macrocarpa 

Benth. oe ° 
Acacia auriculiformis A. Cunn. ex Benth. is Si a 
Acacia basedoww Maid. .. he ee oe a x bn ae 
Acacia aff. bidwillii Benth. ake me ae ie FR nae x 
Acacia brachystachya Benth. 
Acacia calcicola Forde et Ising 
Acacia cambagei R. T. Bak. 
Acacia chisholmii F.™M. Bail. xf <7: & 
Acacia colletioides Benth. a st 2s SF xX <2 wt A 
Acacia conspersa F. Muell. oe ae ah ee 3 Me x 
Acacia coriacea DC. 
Acacia cowleana Tate 4 
Acacia cunninghamw Hook. 
Acacia cuthbertsonii Luehm. : F ne a x 
Acacia cyperophylla F. Muell. ex Benth. a sit x 
Acacia dictyophleba F. Muell. a x 
Acacia difficilis Maid. th oe ee a ke eh fd 
Acacia dimidiata Benth. .. oe af b. ne cn a x 
Acacia dineura F. Muell. oF Be 2 A 
Acacia aff. doratoxylon A. Cunn. ae Bey she x Be ae 
Acacia drepanocarpa F.Muell. .. eG 0 wis Ke a x 


XX *KXX*X: 


xX XK X 
XE OK OK 2S 28 xX 
KK XK 
Ks 


x KS 6 


PROCEEDINGS OF THE LINNEAN SociETy or NEw SourH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 


bo 
ies) 
~ 


Acacia 
Acacia 
Acacia 


dunnvw (Maid.) Turrill 
estrophiolata F. Muell. 
farnesiana (L.) Willd. 


Acacia froggatti Maid. 


Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 
Acacia 


galioides Benth. 

aff. genistoides A. Cunn. ex ; Benth. 
georginae FE. M. Bail. Bye 
gonocarpa F. Muell. : 
gonoclada F. Muell. ex Benth. 
hammondu Maid. 

hemignosta F. Muell. 

hemsleyt Maid. 

hiliana Maid. 

aff. hippuroides Heward ex Benth. 
holosericea A. Cunn. ex G. Don. 
humifusa A. Cunn. ex Benth. 
impressa F. Muell. 

jennerae Maid. 

kempeana F. Muell. 

aff. kempeana F. Muell. 
lamprocarpa O. Schwarz 

latescens Benth. 

latifolia Benth. 

leptocarpa A. Cunn. ex Benth. 
aff. leptocarpa A. Cunn. ex Benth. 
leptophleba F. Muell. tea 
ligulata A. Cunn. ex Benth. 
linarioides Benth. 

linophylla W. V. Fitzg. 
luehmannii F. Muell. : 
lycopoduifolia A. Cunn. ex Hook. 
aff. lycopodiifolia A. Cunn. ex Hook. 
aff. lycopodiifolia A. Cunn. ex Hook. 
aff. lycopodiifolia A. Cunn. ex Hook. 
lysiphloia F. Muell. ex Benth. ~ 
mattlandit F. Muell. Me 
malloclada Maid. et Blakely O06 
microbotrya Benth.. e 
minutifolia F. Muell. 

monticola J. M. Black 
mountfordiae Specht 3 
murrayana F. Muell. ex Benth. 
nuperrima Bak. f. 

oligoneura F. Muell. 

oncinocarpa Benth. 

oswaldi F. Muell. 

aff. oswaldii F. Muell. 5 
pachyphloia W.V. Dee ex Maid. 
pallida F. Muell. 

pellita O. Schwarz 

peuce EF’. Muell. 

phlebocarpa F. Muell. 

plectocarpa A. Cunn. ex Benth. 
polystachya A. Cunn. ex Benth. 
praelongata F. Muell. oor 
proxima Maid. 

pruinocarpa Tindale 

ptychophylla F. Muell. 

pyrifolia DC. 

ramulosa W.V. Fitzg. 

retiventa KF. Muell. 

salicina Lindl. : 

sericata A. Cunn. ex Benth. 
sessiliceps F. Muell. 

shirleyt Maid. : 


PROCEEDINGS OF THE LINNEAN SooreTy oF NEw SoutH WALm#s, VoL. 9 


C.A. 1834 \WARID, IDE we (Et 
O68 x 
x x ane 
x x x 
oe x 
x x 
x 
x Ave 
x x x 
ee x x oe 
DG "4 x >< 
Be x in x 
x x x 06 
eee x Op x 
x x x x 
sive x x 
nie x 
x 
x 
x 
x 
os x 
x x 
Des ae x 
x x Fe 
Se x 
x — 
aad aft Xx 
x x 
5.6 x 
x< : x< 
x x a) ae 
x x x x 
x x or 
bie x 
x 
x ie see 
x x x ee 
att x 
x nee 
yom x 
Xx x 
ae xX x 
x 
x as 
o x x 
see x 
a3 x Xx 
x ns 
xX 
X 
x 
ae Xx 
i's x 
x ye Oa 
as x< x 
x Ke 
x X ee ws 
x Xx x xX 
x ae a: 
he x x 
x a5 a ae 
X xX x 
6, Part 4 


238 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Acacia aff. stbirica S. Moore 33 ws ae? ae x 
Acacia aff. signata F. Muell. ae aoe an ah x Be 
Acacia simsit A. Cunn. ex Benth. sig Al we ay x 
Acacia spondylophylla F.Muell. .. E aie ae x x 
Acacia stenophylla A. Cunn. ex Benth. .. Br ss x x 
Acacia stigmatophylla A. Cunn. ex Benth. ae me HA 
Acacia stipuligera F. Muell. ae ue fd ne x 
Acacia stipulosa F. Muell. we a a9 Zu er, 
Acacia strongylophylla F.Muell. .. un ae a6 x 
Acacia subternata F. Muell. : 5 6 
Acacia sutherlandw F.Muell. 
Acacia tanumbirinensis Maid. 
Acacia tenuissima F. Muell. a a a0 bed x 
Acacia tetragonophylla F. Muell. .. a ats yd x 
x 
x 


a 
KK eX 
x ae 


REN OS 


Acacia torulosa Benth. 

Acacia translucens A. Cunn. ex Hook. ne os i 
Acacia tumida F.Muell. .. : “Ee a wee Me x x 
Acacia umbellata A. Cunn. ex Benth. ae cols x 
Acacia validinervia Maid. et Blakely .. Kis os x 
Acacia victoriae Benth. : 56 ae # aby x 
Acacia wickhamii Benth. .. oe a ae es =f ae x 
Acacia wilhelmiana F. Muell. a ea eh ae sé st 

Acacia wylocarpa A.Cunn. ex Benth. .. Bs se x x 

Acacia yirkallensis Specht. . 

Albizia canescens Benth. = a fs, san 5 58 

+Albizia lebbek Benth. Eee Bis oe: xX ae 
Cathormion umbellatum (Vahl) Kostermans wi ai Ss x 
Dichrostachys cinerea (L.) Wight et Arn. subsp. malesiana 

Brenan et Brummitt . : ai oF ua 
Dichrostachys spicata (F. Muell.) Domin. nf ie i x x 

*Leucaena leucocephala (Lam.) de Wit 

*Mimosa acanthocarpa Poir. 

*Mimosa pigra L. 

*Mimosa pudica L. as are xi ee a ae 
Neptunia dimorphantha Domin a hs x x x 
Neptunia gracilis Benth. forma glandulosa Windler .. nie x x 
Neptunia gracilis Benth. forma gracilis x 
Neptunia major (Benth.) Windler : Bt he ot a Be 
Neptunia monosperma F.Muell. .. on he a x x x 


OS ci OS Oe OS ce cso OX OS OS OK Gs 


XXXKXXX 


IS OS CK OX 6 


CAESALPINIACEAE 
Bauhinia cunninghamii (Benth.) Benth. at we x x XK 
Bauhinia hookeri F. Muell. ae a Se Hite ee Bs 
Cassia absus L. oy: 2 ae Aes a re by oe, x 
*Cassia alata L. 
Cassia artemisioides Gaud. ex DC. ae 
Cassia costata J. F. Bail. et C. T. White 
Cassia desolata F. Muell. 
Cassia glutinosa DC. Me, aes ae a as > Le 
Cassia harneyt Specht ae Ne he he oe a se Se x 
Cassia helmsti Symon AY a ae ote oe x 52 x s3 
Cassia leptoclada Benth. .. “4 ae a Sy aa oe a 4 
Cassia luersseniti Domin .. We ihe a aS x fo a3 
Cassia magnifolia F. Muell. Se fs ate be, ae me x 53 
Cassia mimosoides L. oh a ie ae ot of x x 
Cassia eremophila <A. Cunn. ex Vogel var. coriacea 
(Benth.) Symon : x 
Cassia eremophila A. Cunn. ex Vogel var. eremophila x 
Cassia eremophila A.Cunn. ex Vogel var. platypoda 
(R.Br.) Benth. : ay wa Aes bE x 
Cassia eremophila A. Cunn. ex Vogel var. zygophylla 
(Benth.) Benth. 5% ‘ 
Cassia notabilis F. Muell. 
*Cassia obtusifolia L. 
*Cassia occidentalis L. ; oe ae ia a os ee. a2 
Cassia oligoclada F. Muell. =f de os sf x x x 


xXXXX 


TSS OK IS 
x 


Sax 
Ka 
x 


SOR eS 6 


PROCEEDINGS oF THE LINNEAN Socrety or New Souta Wates, Vou. 96, Part 4 


G. M. CHIPPENDALE 


239 


fo 
P 


V.R.D. D. 


& G. 


Cassia oligophylla EF. Muell. var. oligophylla 
Cassia oligophylla F. Muell. var. sericea Eee 
Cassia phyllodinea R.Br. : 
Cassia planitiicola Domin a 
Cassia pleurocarpa ¥'. Muell. var. ‘pleurocarpa 
Cassia pruinosa F. Muell. : 

Cassia pumila Liam. 

Cassia sturiu R.Br. 5 

*Cassia surattensis Burm. f. 

Cassia timoriensis DC. 

*Cassia tora L. 

Cassia venusta F. Muell. 


Erythrophleum chlorostachyum (F. Muell.) Hennings ex 


Taub. 


Guilandina bonduc Thy, an 
Labichea lanceolata Benth. 
Labichea nitida Benth. 
*Parkinsonia aculeata L. 


Peltophorum pterocarpum (DC.) Backer ex K. Heyne 


Petalostylis labicheoides R.Br. var. cassioides Benth. 


Petalostylis spinescens #. Pritz. 
Piliostigma malabaricum 
(Korth.) de Wit 


(Roxb.) ‘Benth. var. 


*Tamarindus indicus L. 


PAPILIONACEAE 


Abrus precatorius L. 

* Aeschynomene americana be 
Aeschynomene indica L. .. 
Alysicarpus longifolius Wight et ‘Arn. 
Alysicarpus rugosus (Willd.) DC. 


Alysicarpus vaginalis ( 


Dp) Yon 


Atylosia cinerea F. Muell. ex Benth. : 
Atylosia grandifolia F.Muell. ex Benth, 
Atylosia marmorata Benth. Me as 
Atylosia reticulata (Ait.) Benth. .. 
Bossiaea phylloclada F. Muell. 
Brachysema bossiaeoides Benth. 


Brachysema chambersii (EF. Muell. ) F. Muell. ex ‘Benth. 


Burtonia polyzyga (F. Muell.) Benth. 
Burtonia simplicifolia F. Muell. et Tate.. 
Burtonia subulata Benth. .. ‘ 5 


Canavalia 
Canavalia 


cathartica Thouars 
maritima (Aubl.) Thouars 


Clianthus formosus (G. Don) Ford. et Vickery 
*Clitoria ternatea L. 5 a 


*Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 

*Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 
Crotalaria 


acuminata DC. 

ealycina Schrank 5 

crispata F. Muell. ex Benth. 

aff. crispata F. Muell. ex Benth. 
cunningham R.Br. 

dissitiflora Benth. var. dissitiflora 
dissitiflora Benth. var. rugosa Benth. 
tincana L. Ss che : 
juncea L. 

linifolia Lf.  .. 

mitchellia Benth. var. tomentosa Ewart 
nana Burm. f. 


novae-hollandiae DC. var. lasiophylla Benth... 


novae-hollandiae DC. var. novae-hollandiae 
pallida Ait. 

retusa L.. 

strehlowit B. rites 

trifoliastrum Willd. 

verrucosa Li. 


*Cyamopsis tetragonoloba (L.) Taub. 


acidum 


SOK OS OS NK OOK ON OS 


mS OS 6 


KOK | 


De XK OK OK 


x 


SOON DSB 


XXX: 


Xe Ket 


Xe 


SE KEG Ke 


x5 


SN PG 


xx Xs xX X 


xX X 


xX XXX XK XK XK KK XK XK OX 


a PIP GaGa a> GD GsG 


OS 


EIRCOM SOR 


KEDCON, 


PROCEEDINGS OF THE LINNEAN SocretTy or New SoutH WALES, Vou. 96, Part 4 


240 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. & G. 


Cyclocarpa stellaris Afz. ex J.G. Bak. .. M a at va ws x 
Daviesia arthropoda F. Muell. ae a ake “2 x oe a a 
Daviesia reclinata A.Cunn. ex Benth. .. ie ae Bes e% dhs x 
Davwiesia sp. Se ae a ae ae : 
Dendrolobiwm stipatum S. T. Blake Ss as as At Mt 2 x 


Xx 


PROCEEDINGS OF THE LINNEAN Socipty oF New SoutH WALES, Vou. 96, Part 4 


Derris trifoliata Lour. 


Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 
Desmodium 


Dicerma biarticulatum ( 


campylocaulon F. Muell. ex Benth. 

filiforme Zoll. et Morr. 

gangeticum DC. : a Bes ie ae 
mueller? Benth. a we ate as x 
aff. mueller? Benth. ae re ee 
neurocarpum Benth. Be a “e x 
aff. neurocarpum Benth. 

polyneurum 8S. T. Blake 

aff. polyneurum S. T. Blake 

tortuosum (Sw.) DC. 

trichostachyum Benth. 

aff. trichostachyum Benth. 

umbellatum DC. 

L.) DC. 


Dunbaria singuliflora F. Muell. 

Hriosema chinense Vog.  . 

Erythrina variegata L. var. ‘orientalis (L.) Merr. ae ne 
Erythrina vespertilio Benth. ae. ae x 
Galactia megalophylla (¥. Muell.) Ui Fel, Willis 

Galactia muellert Benth. 


Galactia tenuiflora (Klein ex Willd.) Wight et Arn. 
Gastrolobium grandiflorum F. Muell. a oe 
Glycine canescens F. J. Herm. 

Glycine clandestina Wendl. 

Glycine falcata Benth. 


SOS 26 eS 


Glycine tabacina (Lab.) Benth. 


Glycine ton 


ventosa Benth. 


Hovea longifolia R.Br. ex Ait. var. longifolia , 


Hovea long 
Benth. 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 
Indigofera 


Isotropis atropurpurea F. Muell. 
Isotropis parviflora Benth. 

Isotropis wheelert F. Muell. ex Benth. 
Isotropis winneckeana F. Muell. 


Jacksonia 
Jacksonia 
Jacksonia 
Jacksonia 
Jacksonia 
Jacksonia 
Kennedya 


ufoia R.Br. ex Ait. var. pannosa (A. Cunn.) 


Gamedonn E. Tent. : 

brevidens Benth. var. Breaidene 5 
brevidens Benth. var. uncinata Benth. 
colutea (Burm. f.) Merr. 

cordifolia Heyne 

dominii Hichler. . 

george HK. Pritz. 

glandulosa Willd. 

haplophylla F. Muell. 

hirsuta L. 

aff. hirsuta L. 

leucotricha E. Pritz. 

linifolia (L.f.) Retz. 

monophylla DC. a 
parviflora Heyne ex Wight et Arn. 
saxicola KF. Muell. ex Benth. 

subulata Heyne. . 

tinctoria L. F 

trita L.f.. 


SOX XE ea, ORS OK OS 


XS 5 


anomala Ewart et Morr. 
dilatata Benth. .. 

odontoclada F'. Muell. 
ramosissima Benth. 

thesioides A. Cunn. 

vernicosa F. Muell. ex Benth. 
prorepens (F. Muell.) F. Muell. 


BS OK OX S 


x 


x: 


XO 


NE DK OS EK SK OS ER OK OK OK OS OS ORK OE KE OOK 


Oe KOS XE 


So 


xX OK 


GK FE 


G. M. CHIPPENDALE 241 


i 


C.A. Be VekveLD) ae: & G. 


Lotus australis Andr. var. australis 

Lotus australis Andr. var. exstipulatus J M. Black 
Lotus cruentus Court 

Lourea obcordata Desv. e. ee 
+Medicago polymorpha L. var. brevispina (Benth. ) ‘Heyn 
}+Medicago polymorpha L. var. vulgaris (Benth.) Shinners 
}+Medicago truncatula Gaertn. var. truncatula at 
Mirbelia viminalis (A. Cunn.) C. A. Gardn. 

Moghania involucrata (Benth.) Kuntze 

Moghania lineata (Roxb.) Kuntze 

Moghania parviflora (Benth.) Kuntze 

Moghania pauciflora (Benth.) Kuntze 

Mucuna gigantea (Willd.) DC. é 

Mucuna urens (L.) DC. var. papuana F. M. Bail. a 
Paratephrosia lanata (Benth.) Domin .. ee x 
Phaseolus lathyroides L. 5 me a M2 <3 =e as 
{Phaseolus mungo L. hs a oy et Ht ni ae x 
+Phaseolus radiatus L. ee af: aS ne Ud st x 
Pongamia glabra Vent. : 

Pongamia pinnata (L.) Pierre 

Psoralea archert F. Muell. .. be mes nie So a Me ae 
Psoralea badacana (Blanco) Benth. var. badacana a ate ae ate x 
Psoralea badacana (Blanco) Benth. var. grandiflora Benth. x 
Psoralea balsamica F. Muell. : 

Psoralea cinerea Lindl. 
Psoralea eriantha Benth. 
Psoralea leucantha F. Muell. 
Psoralea martini F. Muell. 
Psoralea patens Lindl. AG Se ow a x 
Psoralea plumosa ¥. Muell. ie ats te oie x 
Psoralea pustulata F. Muell. aM me an x x 
Psoralea schultzii F. Muell. a oF 5 Be ae x 
Psoralea walkingtoni F. Muell. 
Pitychosema anomalum F. Muell. 
Ptychosema stipulare J. M. Black 
Ptychosema trifoliolatum F. Muell. 
Pycnospora lutescens (Poir.) Schindl. 
Rhynchosia acutifolia F. Muell. ex Benth. 
Rhynchosia australis Benth. ‘ 4 a ie oe 
Rhynchosia minima (L.) DC. : 3 <a x x 
Rhynchosia rhomboidea F. Muell. ex Benth. 

Rothia trifoliata Pers. 

Sesbania benthamiana Domin ar se a sh ot SN 
Sesbania brachycarpa F.Muell. .. 5 Se on x 
Sesbania cannabina (Retz.) Poir. var. cannabina : x a x x 
Sesbania cannabina (Retz.) Poir. var. sericea (Benth.) 

N. T. Burbidge om 2s a 
Sesbania chippendaler N. T. ‘Burbidge “i fe ae x ~< a 
Sesbania erubescens (Benth.) N.T. Burbidge .. 5% _ oe < 
Sesbania formosa (F. Muell.) N.T. Burbidge .. Si Ss sa, x 
Sesbania javanica Miq. : ae oy 
Sesbania sesban (L.) Merr. Re Be ay: we 
Sesbania simpliciuscula F.Muell. ex Benth. var. 

simpliciuscula .. a oe a a at ae she x 
Smithia conferta Sm. 

*Stylosanthes humilis H.B. K. 


Cx) 


x 


SK ROK S 


0 


Gh OCD Oe DK KK OIG 2 
x 


GS OS ONG 


XG ee 


KGS, 5 OX 


SOK. 


Swainsona beasleyana (F. Muell. yA. ‘Lee subsp. beasleyana x 
Swainsona burkei F.Muell. ex Benth. subsp. acutt- 

carinata A. Lee os x 2% 
Swainsona burkei F. Muell. ex Benth. subsp. burkei .. x x 
Swainsona campylantha F. Muell. : = 
Swainsona canescens (Benth.) F. Muell. var. canescens S< 
Swainsona canescens (Benth.) F. Muell. var. horneana 

J. M. Black E x 
Swainsona cyclocarpa F. Muell. var. cyclocarpa Se Sits 
Swainsona flavicarinata J. M. Black Be oe ac S< x 


PROCEEDINGS OF THE LINNEAN Socrety oF New SoutH Watss, Vou. 96, Part 4 


242 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. &G. 

Swainsona microcalyx J.M. Black subsp. microcalyz .. x 

Swainsona microphylla A. Gray subsp. affinis A. Lee x 

Swainsona microphylla A. Gray subsp. glabrescens A. Lee xX 

Swainsona microphylla A. Gray subsp. pallescens A. Lee x 

Swainsona oligophylla F. Muell. ex Benth. x 

Swainsona oroboides F. Muell. ex Benth. subsp. oroboides x 

Swainsona phacoides Benth. subsp. phacotdes .. x 

Swainsona rigida (Benth.) J.M. Black .. x 

Swainsona unifoliata F. Muell. x 

Swainsona villosa J.M. Black .. »< 56 
Templetonia egena (KF. Muell.) Benth. x eet x 
Templetonia hookert (F. Muell.) Benth. x x x 
Tephrosia brachycarpa F. Muell. ex Benth. x x Bi 
Tephrosia coriacea Benth. fu x x 
Tephrosia eriocarpa Benth. x x x 
Tephrosia filipes Benth. : x a 
Tephrosia flammea ¥F. Muell. ex Benth. : be Bye me x a 
Tephrosia juncea Benth. .. a iis 2 sa ae a x 
Tephrosia leptoclada Benth. : Bis ave are ee 4 x 
Tephrosia macrocarpa Benth. Be me he ae we x 4 a: 
Tephrosia oligophylla Benth. ; ae at us if Se x 
Tephrosia phaeosperma F. Muell. ex Benth. .. 7 x x x x 
Tephrosia polyzyga F.Muell. ex Benth. x a 
Tephrosia porrecta R.Br. ex Benth. is x 
Tephrosia purpurea (L.) Pers. var. axillaris Bak. f. 4 
Tephrosia purpurea (L.) Pers. var. brevidens Benth. .. ae x 
Tephrosia purpurea (li.) Pers. var. longifolia Benth. .. x x x x 
Tephrosia purpurea (L.) Pers. var. purpurea x x she x 
Tephrosia remotiflora F. Muell. ex Benth. a x Xx x x 
Tephrosia rosea F.Muell. ex Benth. es ae oe ist We x at 
Tephrosia sphaerospora F. Muell. ah ae x De 
Tephrosia simplicifolia F. Muell. ex Benth. ee x 
Tephrosia stuartit Benth. x oc 
Tephrosia aff. stuartit Benth. Sid hs se ore x 510 
Tephrosia uniovulata F.Muell. .. ihe Rs te x x Xx 
Trigonella suavissima Lindl. oe a Bie ak x Ss a oe 
Uraria cylindracea Benth. oe eis ne i x Xx 
Uraria lagopodoides (L.) Desv. ex “DC. bs Me x 
+Vicia calcarata Desf. : i on ers x oe 
Vigna canescens C. T. White 5 ae te ee she ae on x 
Vigna lanceolata Benth. var. filiformis Benth. ee ae 33 x x 
Vigna lanceolata Benth. var. latifolia C.T. White  .. x se des oe 
Vigna lanceolata Benth. var. lanceolata 3 se bi x x x 
Vigna luteola (Jacq.) Benth. ie ae - ou: “es a a x 
Vigna marina (Burm. f.) Merr. .. ade bi sit x o¢ 
Vigna vewillata (L.) Benth. var. youngiana F. M. ‘Bail... : hs x 
Zornia albiflora Mohlenbrock Ree ae x .- 
Zornia areolata Mohlenbrock or or =i # aks or x 
Zornia chaetophora F. Muell. at ois fr ae Ae sf x 26 
Zorma gibbosa Span : x 
Zornia muriculata Mohlenbrock x 
Zornia nervata Mohlenbrock x 

GERANIACEAE 

EHrodium aureum Carolin x 

tHrodium botrys (Cav.) Bertol. x< 

tHrodium cicutarium (L.) L’Her. x 

EHrodium crinitum Carolin ue : x 

Hrodium cygnorum Nees ssp. cygnorum x 

Erodium cygnorum Nees ssp. glandulosum Carolin x 

OXALIDACEAE 

Oxualis corniculata L. x 

tOxalis corymbosa DC. x 

tOualis pes-caprae L. x 


PRoOcEEDINGS oF THE LINNEAN SocrETy oF NEw SoutH WatsEs, Vou. 96, Part 4 


G. M. CHIPPENDALE 243 


C.A. Jit, \WalReID IDL eke (Cp 


ERYTHROXYLACEAE 
Erythroxylum ellipticum R.Br... bye a be 8 of bie x 


ZYGOPHYLLACEAE 
Nitraria schobert L. Ai 
Tribulus angustifolius (R. Br.) Benth. 
Tribulus astrocarpus EF. Muell. ; es ee a ne 
Tribulus bicolor F. Muell. ie 5a fe ne an on x a 
Tribulus cistoides L. He Be ae - Be a oe a x 
Tribulus hystrix R.Br. 
Tribulus macrocarpus F. Muell. ex “Benth. 
Tribulus occidentalis R.Br. F ae te se oe 
Tribulus pentandrus Benth. cs ao Sis xo ge x 
Tribulus renunculiflorus F. Muell. 
Tribulus solandri F. Muell. 
Tribulus terrestris L. 
Zygophyllum ammophilum F. Muell. 
Zygophyllum apiculatum F. Muell. 
Zygophyllum aurantiacum Lindl. 
Zygophyllum compressum J. M. Black 
Zygophyllum eremaeum (Diels) Ostenf. 
Zygophyllum glaucescens F. Muell. 
Zygophyllum howittii F. Muell. 
Zygophyllum iodocarpum F. Muell. 
Zygophyllum prismatothecum F. Muell. 
Zygophyllum tesquorum J. M. Black 


xX X 
~N 
x 
x 


xX XX 


dG 


OCS Cle 


Sh OK KOK OK OK ONES OX OS 2K G 


RUTACEAE 
Atalantia recurva Benth. 
Boronia affinis R.Br. ex Benth. ; ae 
Boronia artemisiaefolia F. Muell. var. artemisiaefolia Ae ac Aes " 
Boronia artemisiaefolia F.Muell. var. wilson F.Muell.  .. Be x 
Boronia lanceolata F. Muell. 
Glycosmis trifoliata Spreng. 
Micromelum minutum (Forst. f.) Wight « et Arn. 
Zanthoxzylum parviflorum Benth. AD 


x X X 


XxX XX: 


SIMARUBACEAE 
Brucea javanica (L.) Merr. a a ti F ze oe x< 
Harrisonia browni A. Juss. 
Suriana maritima L. 


x XxX X 


BURSERACEAE 
Canarium australianum F.Muell. var. australianum .. ae ~~ eve x 
Canarium australianum F. Muell. var. glabrum Leenh. oe ne ss x 


MELIACEAE 
Aglaia elaeagnoides (A. Juss.) Benth. : 
Dysoxylum oppositifolium F. Muell. ex DC. 
Melia composita Willd.  .. sh as ie 
Owenia acidula F. Muell. .. a: ae an a x 
Owenia reticulata F. Muell. = a8 ae me x F 
Owenia vernicosa F. Muell. ee os oes oi rae oie x 
Vavaea australiana 8. T. Blake 
Xylocarpus australasicus Ridl. 
Xylocarpus granatum Koenig 


x XX 


LEIK OS OS 


POLYGALACEAE 

Comesperma aphyllum R.Br. ex Benth. . . 

Comesperma secundum Banks ex DC. : 

Comesperma sylvestre Lindl. oi sie i oe x 

Comesperma viscidulum F. Muell. Ss 6 Wee x 

Polygala elongata Klein. .. sic a Si ae 
Polygala eriocephala F. Muell. ex “Benth. ae Ae As an x 
Polygala easquarrosa Adema sae ; oe ete MS 4h aA 
Polygala linariifolia Willd. Fa a m! * os x SX 


x * 


KOS OX) OS 


PROCEEDINGS OF THE LINNEAN SooreTy or New South Wates, Vou. 96, Part 4 


244 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Polygala longifolia Poir. 

Polygala orbicularis Benth. 

Polygala rhinanthoides Sol. ex Benth. 
Polygala stenoclada Benth. 

Polygala teppert F. Muell. .. ts Be ie ae +: : i 
Salomonia oblongifolia DC. a ng ae sue a ee an x 


Mw 
7 


x X X 


SES ek 


EUPHORBIACEAE 

Adriana acerifolia Hook. .. is a Lvs 53 A 
Adriana hookert (F. Muell.) J. Muell... ae sits x BS ae 
Andrachne decaisnei Benth. he oN a oi zh x x 
Antidesma dallachyanum Baill. ; a Be fe ts Ae 
Antidesma ghaesembilla Gaertn. .. st ae a Me ee x 
Antidesma parvifolium F. Muell. 
Antidesma schultzi Benth. 
Beyeria bickertonensis Specht 
Breynia oblongifolia J. Muell. 
Breynia stipitata J. Muell. ay ads =} ae 46 ae 
Bridelia monoica (Lour.) Merr. .. ae 5 ae JE ss x 
Bridelia ovata Dene. aa a sn “is iy a exe AS x 
Calycopeplus sp. A Oy 2 Se: Be sis see ah x 
Croton arnhemicus J. Muell. ae ae Sb: ff ne as x x 
Croton schultz Benth. a 6 ue = 35 ao =i Ae x 
Croton tomentellus F. Muell. ie ch a in ste sins x 

Croton verreauaii Baill. > 
Dissiliaria tricornis Benth. ae a ig bus Fs a A x 
Drypetes australasica (J. Muell.) Pax et Hoffm. Ne fi ae as x 
Huphorbia alsinaefiora Baill. 4d Fre : x 
Euphorbia armstrongiana Boiss. rs a ae a 5 igs x 
Huphorbia atoto Forst.f. .. by te Ff in =f) ay ‘ x 
Huphorbia australis Boiss. 
Huphorbia boophthona C. A. Gardn. 
Euphorbia coghlani F.M. Bail. Be as ae 
Euphorbia comans W.V. Fitzg. .. he ie a ae x 

*Huphorbia dentata Michx. .. te es ed he: a 55 aS 
Euphorbia drummondii Boiss. 5 Be ay te x x x 
Huphorbia eremophila A.Cunn. .. Ne aie Be x x x 
Euphorbia filipes Benth. .. ae as ate a! 
Euphorbia finlaysonii J.M. Black = bie i x 

*Huphorbia heterophylla L. 5 a8 
Huphorbia hirta L. ae ae we Ha se as ae by 
Huphorbia micradenia Boiss. ae i ore fa x x 
Euphorbia mitchelliana Boiss. var. mitchelliana Bs ee x x< 
Huphorbia mitchelliana Boiss. var. stenophylla Benth. 
Euphorbia muelleri Boiss. ve a3 ay sit is 

tHuphorbia peplus L. P a a vie x 
Euphorbia petala Ewart et Kerr. ‘ Bai aes x b3 a 
Euphorbia schizolepis F. Muell. ex Boiss. bd as aie sig x 
Huphorbia schultzii Benth. a se i Ne ee x x 
Huphorbia stevenii F.M. Bail. he ae Ne Bt x x 
Euphorbia tannensis Spreng. he 
Euphorbia vachelii Hook. et Arn. 8s ee ee Sie 
Euphorbia wheelert F.M. Bail. .. at aie en x 
Euphorbia sp. ae ae ae ag x sig sie ae 
Hxcaecaria agallocha rae By ae Om Ss ee A x x 
Excaecaria parvifolia J.Muell. .. Ae Be ss of: x 
Glochidion ferdinandi (J. Muell.) F.M. Bail. var. fer- 

dinandi 
Glochidion ferdinandi (J. Muell.) F. M. Bail. var. _ supra: 
axillaris (Benth.) F. M. Bail. 

Hemicyclia lasiogyna F. Muell. 
Hemicyclia sepiaria Wight et Arn. ek a of ae 

*Jatropha gossypifolia L. .. ae ie ae a x 
Macaranga involucrata (Roxb.) Baill. 
Macaranga tanarius (L.) J. Muell. 
Mallotus nesophilus F. Muell. ex J. Muell. 


x 


SESS SK SSE SEES 


xX X 
x XO DM NOC OMEN ON, De a OS 


XXKXKXKXKXXKXK 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Watss, Vou. 96, Part 4 


G. M. CHIPPENDALE 


245 


V.R.D. D. & G. 


*Micrococca mercurialis (Li.) Benth. : 
Petalostigma banksiz Britten et S. Moore 


Petalostigma haplocladum Pax et K. Hoffm. 
Petalostigma quadriloculare F. Muell. 


Benth. 


var. glabrescens 


Petalostigma quadriloculare F. Muell. var. nigrum Ewart 
et Davies ; 
Petalostigma quadriloculare ‘F. Muell. var. “quadriloculare 


Petalostigma quadriloculare 


F. Muell. var. sericeum 


J. Muell. 


Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 


adam aj. Muell. ; 
baccatus F. Muell. ex Benth. 
carpentariae J. Muell. 
ditassoides J. Muell. .. 


Phyllanthus fuernrohru F. Muell. 


Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 

Domin 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 
Phyllanthus 


huntit Ewart et Davies 

lacunarius F. Muell. on 
maderaspatanus L. var. angustifolius Benth. 
maderaspatanus L. var. maderaspatanus .. 
aff. maderaspatanus Li. 

ochrophyllus Benth. .. ae = 
ramosissimus (F.Muell.) J. Muell. 
rhytidospermus F. Muell. ex J. Muell. : 
simplex Retz. var. ae eee ee Muell. 


simplex “Retz. var. simples - 
aff. sumplex Retz. 

thesioides Benth. 

trachygyne Benth.  . 
trachyspermus F. Muell. 
urinaria L. : 

xerocarpus O. Schwarz 


Poranthera coerulea O. Schwarz 
Poranthera ee eh 
Ricimocarpos sp. 


*Ricinus communis Tt, 
Sebastiania chamelaea ( 


L.) J. Muell. 


Securinega virosa (Roxb. ex Willd.) Pax et Hoffm. .. 


ANACARDIACEAE 


Blepharocarya depauperata Specht 
Blepharocarya involucrigera EF. Muell. 


Buchanania 
Buchanania 
Buchanania 
Semecarpus 


AQUIFOLIACEAE 


angustifolia Roxb. 
arborescens (Bl.) Bl. 
obovata Engl... 
australiensis Eng]. 


Ilex arnhemensis (F. Muell.) Loes. 


CELASTRACEAE 


Denhamia obscura (A. Rich.) Meissn. 
Elaeodendron melanocarpum F. Muell. 
Lophopetalum arnhemicum N. Byrnes 
Maytenus cunninghamui (Hook.) Loes. 


STACKHOUSIACEAE 


Macgregoria racemigera F. Muell. 


Stackhousia 
Stackhousia 
Stackhousia 
Stackhousia 


SAPINDACEAE 
Allophylus serratus (Roxb.) Kurz. 3 
Atalaya hemiglauca (F. Muell.) F. Muell. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutrH WaAtLgEs. VoL. 96, 


intermedia F. M. Bail. 
megaloptera F. Muell. 
muricata Lindl. 
viminea Sm. 


ex Benth. 


KXXEX XS: 


Seo 
Ze 


x 


>. 


x X 


SG 


x 


xX x 


KES: 


Na 


Part 4 


246 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.&G. 


Atalaya salicifolia Bl. 


Atalaya varuifolia F. Muell. ex Benth. 


Cardiospermum halicacabum L.  .. 
Cupaniopsis anacardioides (A. Rich.) 
Diplopeltis stuart F. Muell. 
Distichostemon filamentosus S. Moore 


Radlk. 


Distichostemon hispidulus (Endl.) Baill. © .. .. .. x x 


Dodonaea attenuata A. Cunn. 
Dodonaea lanceolata F. Muell. 
Dodonaea microzyga F. Muell. 
Dodonaea oxyptera F. Muell. 
Dodonaea peduncularis Lindl. var. 
Davies 

Dodonaea petiolaris F. Muell. 
Dodonaea physocarpa F. Muell. 
Dodonaea platyptera F. Muell. 
Dodonaea polyzyga F. Muell. 


5 ON ON OX 
x 
x 


coriacea Ewart et 


Dodonaea viscosa Jacq. var. spatulata (Sm.) Benth. .. x 


Ganophyllum falcatum Bl. 


Harpullia leichhardti F. Muell. ex Benth. 


Heterodendrum floribundum KE. Pritz. 
Heterodendrum oleifolium Desf. 


RHAMNACEAE 
Alphitonia excelsa (Fenzl.) Benth. 


Alphitonia incana (Roxb.) Teijsm. et Binn. ex Kurz. 


Alphitonia aff. moluccana Tiejsm. et Binn. 


Alphitonia petret Braid. et White 
Colubrina asiatica (L.) Brongn. 


Spyridium spathulatum (F. Muell.) F. Muell. ex Benth... x 


Ventilago viminalis Hook. 
Ziziphus mauritiana Lam. 
Ziziphus oenoplia Mill. Be 
Ziziphus quadrilocularis F. Muell. 


VITACEAE 


Ampelocissus acetosa (F. Muell.) Planch. he bys 4; if x 


Ampelocissus sp. 

Cayratia trifolia (L..) ‘Domin 
Cissus reniformis Domin 
Cissus repens Lam. 

Leea brunoniana C. B. Clarke 


ELAEOCARPACEAE 
Elaeocarpus arnhemicus F. Muell.. . 


TILIACEAE 
Corchorus acutangulus Lam. 
Corchorus capsularis L. 
Corchorus echinatus Benth. 
Corchorus eldert F. Muell. 
Corchorus fascicularis Lam. 
Corchorus macropetalus Domin 
Corchorus olitorius L. ate 
Corchorus pascuorum Domin 
Corchorus pumilio R.Br. ex Benth. 
Corchorus rostrisepalus Domin 
Corchorus sericeus Ewart et Davies 
Corchorus sidioides F. Muell. 
Corchorus tridens L. 
Corchorus vermicularis F. Muell. 
Grewia breviflora Benth. .. 
Grewia latifolia F. Muell. ex Benth. 
Grewia multiflora Juss. é 
Grewia orbifolia F. Muell. ex Benth. 
Grewia orientalis Benth. 


PROCEEDINGS OF THE LINNEAN SOCIETY 


XXX: 


5c ON OS ON S 


or New SoutH WALES, Vou. 96, Part 4 


MONON ON NS 


5. ON OS 6 


XXXXXKXK SM OX gee DOS OS OOO 


5 OS OX x 


x! 


KS SF 


x 


G. M. CHIPPENDALE 247 


C.A. B.T. V.B.D. D. &G. 


Grewia retusifolia Kurz. x x 
Grewia scabrella Benth. . 4 be. 
Grewia santhopetala F. Muell. ex Benth x x 
Triumfetta appendiculata F. Muell. ae x“ 
Triumfetta denticulata R.Br. ex Benth. .. = 2 
Triumfetta glaucescens R.Br. ex Benth. ae wt x SG x 
Triumfetta micracantha F. Muell. .. oe be 5 x x x a 
Triumfetta plumigera F. Muell. x x x 
Triumfetta rhomboidea Jacq. Be te Ki we ne x 
Triumfetta winneckeana F. Muell.. ae = ae < 
MALVACEAE 

Abelmoschus ficulneus (L.) Wight et Arn. oe aie x x x 
Abelmoschus moschatus Medic. ssp. tuberosus (Span. ) 

Borss. .. ee Be a oe og x 
Abutilon andrewsianum W. V. Fitze. oy: Se ake a es >< x 
Abutilon crispum G. Don .. fe ae x x 
Abutilon cryptopetalum (F. Muell.) F. Muell. ex - Benth. x ie ee 
Abutilon cunninghami Benth. .. : He: a ve > 
Abutilon frasert (Hook.) Hook. ex Walt. ee sé 
Abutilon indicum (L.) Sweet var. australiense Hockr. ex 

J. Britt. 4 £3 ue a of as ae a. an x 
Abutilon aff. indicum (L.) Sweet ats >< a3 4 
Abutilon leucopetalum (KF. Muell.) F. Muell. ex Benth. x x x 
Abutilon macrum F. Muell. : es x a 
Abutilon malvifolium (Benth.) J. M. Black x sy 
Abutilon otocarpum F. Muell. as ae a x x x x 
Alyogyne pinonianus (Gaud.) Fryxell var. pinonianus x Br iy be 
Camptostemon schultzii Mast. ah fs 54 ae a Be x 
Gossypium australe F. Muell. 1 are aa ae > x x >< 
Gossypium bickiti Prokh. .. i bk at a. x x x 
Gossypium cunninghami Todaro .. ee ss ae ae < 
*Gossypium hirsutum L. var. Re (Schum. ) J. B. 

Hutch. .. at a Wee Ae x 
Gossypium sturtianum 5. TEL. ‘Willis var. sturtianum off x Bis - 
Hibiscus brachychlaenus F. Muell. a a ale x x x x 
Hibiscus brachysiphonius F. Muell. x i 
Hibiscus caesius Garcke .. of 348 as = x x x 
Hibiscus crassicalyx J. M. Black . ae - a x es ae ss 
Hibiscus divaricatus Grah. My ie 5% Les ae 3 7 x 
Hibiscus drummondit Turcz. me & ae ts x 
Hibiscus farraget F. Muell. aes te i 58 x ae oa ble 
Hibiscus heterophyllus Vent. a oie ae ate ate ad x 
Hibiscus krichauffianus F. Muell. var. chippendaler 

Fryxell .. oa xx x 
Hibiscus krichauffianus F. Muell. ‘var. brichaufianus 50 x ies ot 
Hibiscus leptocladus Benth. : : ae x x 
Hibiscus menzelii F. D. Wilson and N. Byres a ot a x 
Hibiscus meraukensis Hochr. 5S . a x x x x 
Hibiscus panduriformis Burm. f. .. x x 
Hibiscus pentaphyllus F. Muell. x x 
*Hibiscus sabdariffa L. : x x 
Hibiscus setulosus F. Muell. ae mA aes ic be a ae x 
Hibiscus sturtit Hook. var. campylochlamys Benth. .. aA x< x x 
Hibiscus sturtii Hook. var. grandiflorus Benth. x x b>. a 
Hibiscus sturtti Hook. var. muellert Benth. x see x 
Hibiscus sturtii Hook. var. platychlamys Benth. X x x 
Hibiscus sturtit Hook. var. sturtiz if ee x x 
Hibiscus sturtii Hook. var. truncatus Fryxell .. xe x as a nee 
Hibiscus symonii F. D. Wilson and N. Byrnes Ne st ae sh x 
Hibiscus tiliaceus L. ssp. tiliaceus Sis ae 36 ot x ae x 
Hibiscus trionum L. ae a oa Fa a Ss x x 
Hibiscus zonatus F. Muell... we ar aa aie oe oe oe x 
Lavatera plebeia Sims RY on oe we Se 7 ae Sa 
*Malachra fasciata Jacq. .. ast ft St 8 ne ac ae x 
*Malva parviflora L. i et oa ae ar x 


PROCEEDINGS OF THE LINNEAN Socrety oF New SourH WateEs, Vou. 96, Part 4 


248 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. & G. 


Malvastrum americanum (L.) Torr. sie dies Be x x x 
Plagianthus glomeraius Fie Benth. .. : we x Be ih Me 
*Sida acuta Burm. f. ay + AM ee x x 
Sida ammophila F. Muell. ex J. H. Willis x rsh 

Sida calyxhymenia J.Gay ex DC. x x x 

Sida cardiophylla F. Muell. x x ap 

Sida cleisocalyx F. Muell. x x x 


*Sida cordifolia L. .. a a ap far si x 
Sida corrugata Lindl. var. “angustifolia Benth. 
Sida corrugata Lindl. var. corrugata 
Sida cryphiopetala F. Muell. 

Sida cunninghamu White .. 

Sida fibulifera Lindl. 

Sida aff. fibulifera Lindl. 

Sida filiformis A.Cunn.  .. Bt ae 
Sida goniocarpa (F. Muell. ex Benth. Domin .. 
Sida intricata F. Muell. a's : 
Sida lepida F. Muell. otis ae 

Sida macropoda F. Muell. ex Benth. ae 5 x 
Sida petrophila F.Muell. .. eS Me bs a": ae x* x 
Sida platycalyx F. Muell. ex Benth. x 

Sida rhombifolia L. var. incana Benth. a ue M, e 
Sida rhombifolia L. var. rhombifolia ot ue ae bo * oe x 
Sida rohlenae Domin Bs ze! os AY, f x x i ae 
Sida spinosa L. re ah Se nts x x x 
Sida subspicata F. Muell. ex Benth. ‘y ae A ae fe x 
Sida trichopoda F.Muell. .. ate 5 x x 
Sida virgata Hook. var. phacotricha (F. Muell. ) Benth... x ae a 
Sida virgata Hook. var. eae oe At a a5 x in 4 
Sida sp. : an Xx 
Thespesia lampas (Cav.) Dalz. ex Dells ‘et Gibbs var. 

thespidioides (R.Br. ex Benth.) Fryxell 
Thespesia populnea (L.) Sol. ex Corr. 
Urena lobata L. : a x A 
Urena spencert (Ewart) Fryxell .. 


XX XK XX 
xX 
Bee Grd 
x 


OKO 
XX X 


> ON ON 2S OSS 
x 


x xX 


a ae 


xX xX XK X 


BOMBACACEAE 
Bombax ceiba L. var. leiocarpum Robyns 


x 


STERCULIACEAE 

Brachychiton diversifolium R.Br. .. ae ae a My x x x 
Brachychiton gregorit F.Muell. .. ae = pee x hs oh ote 
Brachychiton paradoxum Schott. .. ee ae By! ff o< Xx Xx 
Brachychiton tuberculata W.V.Fitzg. .. AK a *, Hs Xx 
‘Commersonia crispa Turcz. oe : 

Commersonia melanopetala F. Muell. ae “4 a zt bg 
Dicarpidium monoicum F. Muell. os Bi se af Lt ne x 


x Xx 


Gilesia biniflora F. Muell. x 
Hannafordia bissellia F. Muell. hs ae ae x aN Be mE 
Helicteres cana (Schott.) Benth. ae ae oi 4 Xx 
Helicteres dentata F. Muell. ex Benth. var. dentata ae a he x 
Helicteres dentata F.Muell. ex Benth. var. flagellaris 

Benth. .. i Pe 2 x 
Helicteres dentata F. Muell. ex Benth. var. procumbens 

Benth. .. si Se ne oF MM as st uf 4 x 
Helicteres isora L. .. Sy ae Sh oie 23 ay ae Xx 
Keraudrenia integrifolia Steud. pas we aN "3 x x x 
Keraudrenia nephrosperma (F.Muell.) F.Muell. ex 

J. M. Black ae me ot A Pak x ae x 5 
Melhania oblongifolia F. Muell. = 3 af ag < x Xx Xx 
Melhania ovata (Cav.) Spreng. .. ae oh MN jie x oh ya 
Melochia corchorifolia L. ns ste He an oy ae x 4 
Melochia pyramidata L. .. oe os oe St ei Ae x x 

*Pentapetes phoenicea L. ae ae Be Sis is x 

Rulingia kempeana (F. Muell. ) F. Muell. ex J.M. Black x Ry: fe 
Rulingia loxophylla F. Muell. x He ae x 4 5 


PROCEEDINGS OF THE LINNEAN SocrETyY oF NEw SoutH WatsEs, Vow. 96, Part 4 


G. M. CHIPPENDALE 


Rulingia magniflora F. Muell. 
Sterculia quadrifida R.Br. 

Waltheria indica L. 

Waltheria virgata Ewart et ‘Cookson 


DILLENIACEAE 
Dillenia alata (R.Br. ex DC.) Martelli .. 
Hibbertia browniz Benth. Lie 
Hibbertia cistifolia R.Br. 
Hibbertia dealbata Benth. 
Hibbertia glaberrima F. Muell. 
Hibbertia holizec F. Muell. 
Hibbertia lepidota R.Br. . 
Hibbertia muellert Benth. 


Hibbertia oblongata R.Br. var. brevifolia Benth. 


Hibbertia oblongata R.Br. var. oblongata 
Hibbertia scabra R.Br. ex Benth. 
Hibbertia tomentosa R.Br. .. as 
Pachynema complanatum R.Br. 

Pachynema dilatatum Benth. 

Pachynema junceum Benth. : 
Pachynema sphenandrum F. Muell. “et Tate 


GUTTIFERAE 
Calophyllum australianum F. Muell. 
Calophyllum inophyllum L. 
Calophyllum lonchophyllum_O. Schwarz 
Calophyllum ramiflorum O. Schwarz 
Hypericum gramineum Forst. f. 
Hypericum japonicum Thunb. 


ELATINACEAE 
Bergia ammannioides Roth. a aN 
Bergia pedicellaris F. Muell. ex Benth. 


Bergia perennis (F. Muell.) F. Muell. ex Benth. 


Bergia pusilla Benth. 
Bergia trimera Fisch. et Mey. 
Elatine gratioloides A. Cunn. 


FRANKENIACEAE 
Frankenia connata Sprague 
Frankenia cordata J. M. Black 
Frankenia gracilis Summerh. 
Frankenia muscosa J. M. Black 
Frankenia planifolia Sprague et Summerh. 
Frankenia serpyllifolia Lindl. 
Frankenia speciosa Summerh. 
Frankenia stuartii Summerh. 


CocHLOSPERMACEAE 
Cochlospermum frasert Planch. 
Cochlospermum gillivrayt Benth. .. 
Cochlospermum gregorti F. Muell. .. 


VIOLACEAE 


Hybanthus enneaspermus (L.) F. Muell. var. banksianus 


Domin 


Hybanthus enneaspermus (ie ) ie Muell. var. enneaspermus 


PASSIFLORACEAE 
Adenia australis (R.Br. ex DC.) Engl. 
*Passifiora foetida L. a: ei: 
*Passiflora suberosa L. 


CACTACEAE 
*Opuntia stricta Haw. 


PROCEEDINGS OF THE LINNEAN Society oF NEw Soutru WaAtes. Vou. 96, 


an 


A 


DK OL OK OK OK OK OK OK 


SSSR 


x 


Part 4 


250 NORTHERN THRRITORY PLANTS 


C.A. B.T. 


V.R.D. D. & G. 


THYMELAEACEAE 
Phaleria blumei Benth. var. latifolia Benth. .. ae a Sh ae x 
Pimelea ammocharis F. Muell. aa : at a x x x 
Pimelea microcephala R.Br. oe Ee te site x Bi He Se 
Pimelea punicea R.Br. the a ae ae us sue x SK x 
Pimelea trichostachya Lindl. oe af er ae x 


LYTHRACEAE 


Ammannia 
Ammannia 
Ammannia 
Ammannia 
Ammannia 
Ammannia 


auriculata Willd. 

crintpes F. Muell. 

andica Lam. .. 

multiflora Roxb. 

occultiflora Koehne var. leichhardtii ‘Koehne 
triflora R.Br. ex Benth. 


SEE OX G 


er 


Lawsonia alba Lam. : ae “is 2 ts 5¥3 ne 
Lythrum arnhemicum F. Muell. .. a su No ba anh x 
Pemphis acidula FYorst. et f. : ons es oe i. ae fe 
Rotala diandra (F. Muell.) Koehne os on He x ahh 4 
Rotala mexicana Cham. et Schlecht. ae wee ab sive x ox 
Rotala roxburghiana cere oe ae ae cee is x x 
Rotala verticillaris L. 2 ae a ae aie x 

Rotala sp. 


XXXKXKXXKXKXXKXKX: 


xs 


SONNERATIACEAE 
Sonneratia caseolaris (L.) Engl. .. oo i me m 3 +4 x 
BARRINGTONIACEAE 
Barringtonia acutangula (L.) Gaertn. ssp. acutangula er Be x 
Planchonia careya (F.Muell.) R. Knuth.. = ‘ 


xX 


RHIZOPHORACEAE 
Bruguiera gymnorrhiza Lam. 
Bruguiera parviflora (Roxb.) Wight et Arn. 
Bruguiera rheediw Bl. 
Bruguiera sexangula (Lour. ) Poir. ae ae oe gt Ss 
Carallia brachiata (Lour.) Merr. .. pe te We ee a x 
Ceriops tagal (Perr.) C. B. Robinson er St a ag at x 
Rhizophora mucronata Lam. 2 
Rhizophora stylosa Griff. 


XxxXXKXXKXKXX 


CoMBRETACEAE 
LIumnitzera littorea (Sack) Voigt 
Lumnitzera racemosa Willd. or a an Sf 5H ss 
Macropteranthes kekwickit F. Muell. ae 24 cu 0 Xx x 


x X 


Terminalia 
Terminalia 
Terminalia 
Terminalia 
Terminalia 
Terminalia 


aridicola Domin 

arostrata Kwart et Davies 
bursarina F.Muell. .. 
canescens (DC.) Radlk. 
carpentariae White 
erythrocarpa F. Muell. 


Terminalia ferdinandiana Exell 


Terminalia 
Terminalia 
Terminalia 
Terminalia 
Terminalia 
Terminalia 
Terminalia 
Terminalia 


MYRTACEAE 


grandiflora Benth. 
latipes Benth. .. a 
melanocarpa F. Muell. 
platyphylla F.Muell. .. 
platyptera F. Muell. 
pterocarya EF. Muell. 
sericocarpa EF. Muell. 
volucris R.Br. ex Benth. 


Acmena smithii (Poir.) Merr. et Perry .. 
Baeckea intratropica (F.Muell.) Niedenzu 
Baeckea polystemona F. Muell. 

Baeckea virgata Andr. 


XX: 


x 


PROCEEDINGS OF THE LINNEAN SocieTY or New SoutH Watss, Vou. 96, Part 4 


MOXSKKS KR KG 


XK OK OK KOK XK 


5 oN eX 


G. M. CHIPPENDALE 251 


C.A. IsjfbG (WilbtID5 ID. ae (Gi 


Callistemon aff. viminalis ay ex Gaertn.) G. Don ex 


Loud. 


Calytriz achaeta F. Muell. 

Calytriz arborescens ¥. Muell. 
Calytriz brachychaeta F. Muell. 
Calytriz laricina R.Br. ex Benth. 
Calytriz longiflora F. Muell. 
Calytrix microphylla A. Cunn. 


Cleistocalyz operculata (Roxb.) Merr. et Perry 


Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Eucalyptus 
Hucalyptus 
Eucalyptus 


abbreviata Blakely et Jacobs 
alba Reinw. ex Bl. .. 
apodophylla Blakely et Jacobs 
argullacea W. V. Fitzg. 

aspera EF’. Muell. 

bigalerita F. Muell. 

bleeser? Blakely 

brachyandra F. Muell. 
brevifolia F. Muell. 
camaldulensis Dehnh. j 
clavigera A. Cunn. ex Schau. 
cliftoniana W. V. Fitzg. 
confertiflora KF. Muell. 
cyanoclada Blakely 
dichromophloia F. Muell. 


FHucalyptus ferruginea Schau. 
Hucalyptus foelscheana F. Muell. 


Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 


gamophylla F. Muell. 
gongylocarpa Blakely 
grandifolia R.Br. ex Benth.. 
herbertiana Maid. ‘ 
intertexta R.T. Bak. .. 


Eucalyptus jacobsiana Blakely 


Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Eucalyptus 
Hucalyptus 
Hucalyptus 
Eucalyptus 
Hucalyptus 
Hucalyptus 
Eucalyptus 
Hucalyptus 
Hucalyptus 
Eucalyptus 
Hucalyptus 
Eucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 
Hucalyptus 


jensenit Maid. .. 

latifolia F. Muell. 
mannensis Boomsma .. 
microtheca EF. Muell. 

minmata A. Cunn. ex Schau. 
morrisit R. T. Bak. 
nesophila Blakely 


normantonensis Maid. et Cambage ae 


odontocarpa F. Muell. 
oligantha Schau. 
oxymitra Blakely 
pachyphylla F. Muell. 
papuana F. Muell. 
patellaris F. Muell. 
phoenicea F. Muell. 
polycarpa F. Muell. 
porrecta S. T. Blake 
pruimosa Schau. 
ptychocarpa F. Muell. 
sessilis (Maid.) Blakely 
setosa Schau. stg 
tectifica F. Muell. 
terminalis F. Muell. 
tetrodonta F. Muell. 
thozetiana F. Muell. oo 
aff. transcontinentalis Maid. . . 
trivalvis Blakely 
umbrawarrensis Maid. 
websterana Maid. 


Hugenia armstrong Benth. 
Hugenia bleesert O. Schwarz 
Hugenia eucalyptoides F. Muell. 
Hugenia suborbicularis Benth. 
Fenzlia retusa Endl. 


PROCEEDINGS OF THE LINNEAN Socrety or New SourH Wates, Vou. 96, Part 4 


~ ox 


5 XxX Ke 


0 ON Mo 


5 oN eG 


eee ate 


ee IO) ON rss OGRE Se BOGGS Oe 


6 ON OK ING 


x 


X 


XxXXXXXX: 


5 OS ON OS OS OG ON OK SG KORO K KOK No MN ORS 


<< OS 6 


MOOS OK KK 5 ON 


5 oA os PSO 


252 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Homalocalyx ericaeus F. Muell. 


Leptospermum abnorme F. Muell. ex Benth. 


Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 
Melaleuca 


acacioides F. Muell. 
alsophila A. Cunn. ex Benth. 
argentea W. V. Fitzg. 
bracteata F. Muell. 
cajuputt Powell. . 

dealbata 8. T. Blake 
dissitiflora F. Muell. 
glomerata F. Muell. 
lasiandra F. Muell. 
leucadendron (L.) L. 
linariifolia Sm. 

linophylla F. Muell. 
magnifica Specht 
minutifolia F. Muell. 
nervosa (Lindl.) Cheel 
symphyocarpa F. Muell. 
viridiflora Sol. ex Gaertn. 
Je 


SC O8 OS 5 


SOK 


x xX X 


OSCR OG 


Metrosideros eucalyptoides (F. ae F. Muell. a oe ahs x 
Micromyrtus flaviflora (F. Muell. ) F. Muell. ex J. M. 

Black .. : ats x 
Osbornia octodonta 190 Muell. died as “4 aus 
Thryptomene maisonneurvtt F. Muell. ee ine ic x 
Thryptomene parviflora (F. Muell.) Domin Sh ai x ae aH 
Tristania grandiflora Cheel ie ai te oe bis x x 
Tristania lactiflua F. Muell. : 
Tristania suaveolens Sm. var. glabrescens 1, M. Bail. Ss <3: ri 
Tristania suaveolens Sm. var. suaveolens ae ae abe ae x 
Verticordia cunninghami Schau. . . is we i 5 Be x 
Verticordia sp. ae oa fel it sis “ 
Xanthostemon par adoxus F. Muell. as a: or ae se x 


XS 6 


XE OK KOK OK SOK 


MELASTOMATACEAE 
Melastoma denticulatum Labill. 
Memecylon pauciflorum Bl. a Be ae Ss 2h ne 
Osbeckia australiana Naud. ee, hs Pi an ay iy x 


x XX 


ONAGRACEAE 
Ludwigia adscendens (L.) Hara .. ae ae ato R uss # x 
Ludwigia hyssopifolia (G. Don) Exell .. a “ee a ys x 
Ludwigia octovalvis (Jacq.) Raven ssp. octovalvis ; ae *< x x 
Ludwigia octovalvis (Jacq.) Raven ssp. sessiliflora (Mich. ) 
Raven ae 
Ludwigia perennis ih, 


x X 
x! 
x 


HALORAGACEAE 
Haloragis acanthocarpa Brongn. .. eis Ss ae 3% os < x 
Haloragis glabrescens C.'T. White Ag Sue sts ” x 
Haloragis gossei ¥. Muell. ae a 
Haloragis heterophylla Brongn. var. glaucifolia Schindler .. aD x 
Haloragis heterophylla Brongn. var. Hes ce 
Haloragis odontocarpa F. Muell. 

Loudonia roet (Endl.) Schldl. 
Myriophyllum diococcum F. Muell. 
Myriophyllum verrucosum Lindl. .. 


MRA OX 


6 
x 
28 OS 5 


ARALIACEAE 
Mackinlaya macrosciadea (F.Muell.) F. Muell. .. nt if dig a x 
Schefflera actinophylla (Endl.) Harms .. oe: a a i ie x 


UMBELLIFERAE 
Actinotus schwarzii F. Muell. a: tek ae x 
tApium leptophyllum (DC.) F. Muell. : : x 
Daucus glochidiatus (Labill.) Fisch., Mey. Gi Acre: Tae Sg 


PROCEEDINGS OF THE LINNEAN Society or New Soura WaAteEs, Vou. 96, Part 


G. M. CHIPPENDALE 


Hydrocotyle 


trachycarpa FE. Muell. 


Platysace arnhemica Specht 


Trachymene 
Trachymene 
Trachymene 
Trachymene 
Trachymene 
Trachymene 
Trachymene 


didiscoides (F. Muell. ) ic L. Burtt 
dusenit (Domin) B. L. Burtt A 
gilleniae (Tate ex Domin) B. L. Burtt 
glaucifolia (F. Muell.) Benth. 
microcephala (Vent.) B. L. Burtt 
setosa (O. Schwarz) B. L. Burtt 

villosa (F. Muell.) Benth. 


EPACRIDACEAE 
Leucopogon sp. 


MyRsINACEAE 
Aegiceras corniculatum (L.) Blanco 


PRIMULACEAE 
tAnagallis foemina Mill. 
*Anagallis pumila Swartz 
Samolus valerandi L. 


PLUMBAGINACEAE 
Aegialitis annulata R.Br. 
Plumbago zeylanica lL. 


SAPOTACEAE 
Mimusops elengi L. 
Planchonella arnhemica (F. Muell. ex Benth. ) van in Royen 
Planchonella crocodiliensis van Royen : 
Planchonella laurifolia (Richard) Pierre 
Planchonella sericea (Ait.) Baehni 


EBENACEAE 
Diospyros calycantha O. Schwarz 
Diospyros ferrea (Willd.) Bakh. var 
Bakh. .. we ee 
Diospyros maritima BI. es 
Diespyros montana Roxb. var. timorensis Bakh. 
Diospyros sp. 


. humilis (R.Br.) 


OLEACEAE 


Jasminum 
Jasminum 
Jasminum 
Jasminum 
Jasminum 
Jasminum 
Jasminum 


aemulum R.Br. 
calcarium F. Muell. 
didymum Forst. f. 
lineare R.Br. 

molle R.Br. 2 
simplicifolium Forst. f. 
volubile Jacq. 


LOGANIACEAE 


Fagraea racemosa Jack 


Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 
Mitrasacme 


alsinoides R.Br. ; ae 
arnhemica Specht var. arnhemica .. 
arnhemica Specht var. Ns waar 
connata R.Br. 

elata R.Br. 

easerta K. Muell. 

gentianea F. Muell. 

indica Wight 

laevis Benth. 

laricifolia R.Br. 

longifiora F. Muell. ex "Benth. 

lutea F. Muell. 

multicaulis R.Br. 

nummularia S. Moore 

polymorpha R.Br. 


oso 


KKK OE KK 


ae 


Sia 


Oe 


xX X 


XXX XX 


xXx: 


eX OS KX XOX 


ea sae 


Xx 


PROCEEDINGS OF THE LINNEAN SocreTy oF New SoutH WaAtEs, Vou. 96, Part 4 


254 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. & G. 


Mitrasacme prolifera R.Br. 

Mitrasacme stellata R.Br. var. latifolia Benth... 
Mitrasacme tenuiflora Benth. : ee bs by a a 
Mitrasacme volubilis O. Schwarz as A d. et ay ays x 
Strychnos lucida R.Br. aes she a Eve ate aH x x 


xX X X 
x 


GENTIANACEAE 
Centaurium spicatum (L.) Fritsch a3 bh EAs x bad x x 
Canscora diffusa R.Br. ee ae es a a be a x 


MENYANTHACEAE 
Nymphoides crenatum (F. Muell.) Kuntze is at 53 x 
Nymphoides geminatum (R.Br.) Kuntze ae 50 x sce 
Nymphoides furculaefolia Specht .. site a 
Nymphoides indica (L.) Kuntze xs 
Nymphoides minimum (F. Muell.) Kuntze 


36 OS SK OK OK 


APOCYNACEAE 

Alstonia actinophylla (A. Cunn.) K.Schum. .. a oe y a 
Alstonia ophiowyloides F.Muell. .. a ats BG aE he x 
Alyxia spicata R.Br. ae a a sh ane a si 
Carissa lanceolata R.Br. .. 53 As ™ x x x 
Ervatamia orientalis (R.Br.) Domin 

Ervatamia pubescens (R.Br.) Domin wn ne ae ay af 
Parsonsia velutina R.Br. .. ie SF: “e ee x 
Wrightia pubescens R.Br. ssp. pubescens sas ae se ee ce 
Wrightia saligna (R.Br.) F. Muell. ex Benth. .. 3 ae se x 


x XK XK XK K XK XK XK 


ASCLEPIADACEAE 

*Calotropis procera (Willd.) R.Br. ex Ait. on LA Ae “3 a 
Cynanchum carnosum (R.Br.) Domin .. 6 a a 31 x 
Cynanchum floribundum R.Br... ae sae als x de <i 
Cynanchum pedunculatum R.Br. .. oe ays ee ses x x 
Cynanchum puberulum F. Muell. ex Benth. 
Gymnanthera nitida R.Br. oa aya 5 a x 
Gymnema geminatum R.Br. ae om ae ae tie ae 5/3 x 
Gymnema muellert Benth. 
Gymnema stenophyllum A. Gray be we Sad Ht: of x 
Gymnema trinerve R.Br... ths ae Si ee zit - o6 x 
Hoya sp. am Se ci sa bs x 
Leichhardtia australis R.Br. sie He ue an x 
Marsdenia cinerascens R.Br. ne ne Ss = ae RS x 
Marsdenia velutina R.Br. .. an of oe pes ns a we x 
Marsdenia viridiflora R.Br. k x 
Microstemma glabriflorum F. Muell. a ae ne ve 
Pentatropis linearis Dene. .. es Sa af x 
Pentatropis quinquepartita Benth. O90 C0 oad a 
Sarcostemma australe R.Br. ; es Ne 54 Xx 
Tylophora erecta F. Muell. ex Benth. . ; ke ae si a ot 
Tylophora flecuosa R.Br. .. Ss a oe ay uf ats x 
Tylophora macrophylla Benth. 

Vincetowicum ovatum Benth. 


x SoS 6 OS OS 
xX X 


x OS 6 


MS KOK KK S 


CONVOLVULACEAE 

Bonamia linearis (R.Br.) Hall. f. Sui is bas x x x 
Bonamia pannosa (R.Br.) Hall. f. as 56 es i x x 
Bonamia rosea (F.Muell.) Hall. f. Se ih ve S< 

Convolvulus erubescens Sims xe a ne Be S< 5 wi 
Oressa australis R.Br. ns as ie ae ats one aa x 
Cressa cretica Li. : af ate 7 aa ta: Bn ae S< 
Cuscuta australis R. Br. ae ay ae sy ath x oy F 
Dichondra repens Forst. et f. ate Bt a a hs Ne 90 
Hvolvulus alsinoides L. var. alsinoides 40 ee ae a a 
EHvolvulus alsinoides L. var. decumbens (R.Br.) v. Ooststr. x x x 
EHvolvulus alsinoides L. var. sericeus Benth. = oe ahs ae 
Hvolwulus alsinoides L. var. villosicalyx v. Ooststr. .. x x x 


x xX 


OK OS 2% OX 2S OK OSB 


PROCEEDINGS OF THE LINNEAN Soctety oF NEw SoutH Watss, Vou. 96, Part 4 


G. M. CHIPPENDALE 255 


C.A. B.T. V.R.D. D. & G. 


Ipomoea abrupta R.Br. ang a sea aie x oft NS * 
Ipomoea alata R.Br. : ae ov Di a a g3 
Ipomoea aquatica Forsk. .. ot x Sys 36 x a x x 
Ipomoea aff. aquatica Forsk. ss Die 7 be ai x 2 . 
Ipomoea aff. aquatica Forsk. ae ne ae an ke ae Re y 
Ipomoea aff. biloba Forsk. 
Ipomoea coptica (L.) Roth. ex R. et S. a an 
Ipomoea costata F. Muell. ex Benth. aig Sy Ne x hee 
Ipomoea aff. costata F. Muell. ex Benth. “ee sie oe oh me x 
Ipomoea davenportit F. Muell. as Ge ne 30 x Hy 
Ipomoea eriocarpa R.Br. .. ie a ne aie = x 
Ipomoea flava F. Muell. ex Benth. 
Ipomoea gracilis R.Br. : 
Ipomoea graminea R.Br. 
Ipomoea hederacea Jacq. 
Ipomoea incisa R.Br. ‘ i ne ie Bs 5 
Ipomoea lonchophylla J. M. Black ay tc ae x x 
Ipomoea longiflora R.Br. é : a oe a 
Ipomoea muellerti Benth. .. ue He i an x 
Ipomoea nil (L.) Roth. g ae oe 
Ipomoea brasiliensis (L.) G. F. W. Mey. 
Ipomoea plebecra R.Br. as is a8 ae ey i a bs 
Ipomoea polymorpha R. et S. 4 oe Re < x x x 
Ipomoea racemigera F. Muell. et Tate x 
Ipomoea triloba L. ‘ 
Ipomoea velutina R.Br. 
Jacquemontia browniana v. Ooststr. 
Jacquemontia aff. browniana v. Ooststr. i ap 
Jacquemontia paniculata (Burm. f.) Hall. f. var. paniculata of : x 
Jacquemontia paniculata (Burm. f.) Hall. f. var. tomentosa 

(Warb.) v. Ooststr. 
Merremia aegyptia (L.) Urban 
*Merremia dissecta (Jacq.) Hall. f. 
Merremia gemella (Burm. f.) Hall. f. 5 We ah 
Merremia quinata (R.Br.) v. Ooststr. ... ae 
Merremia tridentata (L.) Hall. f. au) angustifolia ( (J acq. ) 

v. Ooststr. a ae: ay x 
Merremia tridentata (L.) Hall. f. ssp. ‘hastata (Desf.) 

v. Ooststr. ss : 58 
Merremia tridentata (L.) Hall. f. ssp. tridentata 
Operculina aff. brown v. Ooststr. 

Polymeria ambigua R.Br. . 
Polymeria longifolia Lindl. ae 5 22 ie ae fo 
Polymeria pusilla R.Br... st =i m2 Ae ae 2: os x 


xX<ha xe 
xX 
xX 


x 
x 5 ONO ON OW OS ONG 
SS SS OK OS OK ROK OS eK OK og OS OS OSE GS Suki oe SONOS OSSD 


xX xX 


OS XG 
x 
x 


BORAGINACEAE 

+Buglossoides arvense (L.) I. M. Johnston wi ii x et V2 u: 
Coldenia procumbens Ll... ae me ne ee a! as x x 
Cordia subcordata Lam. .. bes 28 a x 
Cynoglossum australe R.Br. var.  drummondii (Benth, ) 

Brand. 

*Hchium lycopsis L. =a 
Halgania cyanea Lindl. var. cyanea a ote tee 
Halgania cyanea Lindl. var. preissiana (Lehm.) Maid. 

et Betche ; 3s a Se Se x 

Halgania erecta Kwart et Rees re 5 me = x 
Halgania glabra J. M. Black a at 6 ae; SK Ar ae 
Halgania solanacea F. Muell. a ar Sg be: x md Se 
Halgania aff. solanacea F. Muell.. . Ag 3 Me 3: aE x 
Heliotropium asperrimum R.Br. sie se Le x as 
Heliotropium ball Domin. a ai a be a2 x 
Heliotropium bacciferum Forsk. .. : oe x aM ah a8 
Heliotropium bracteatum R.Br. var. bracteatum see oh a x 
Heliotropium bracteatum R.Br. var. Renee Benth. .. eh; ee S< 
Heliotropium carpentariae Specht. . SrA ee a et x 
Heliotropium conocarpum F. Muell. ex Benth. ae ete mn x 


x X X 


PROCEEDINGS OF THE LINNEAN SocretTy oF NEw SoutH Wates. Vou. 96, Part 4 


256 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Heliotropium curassavicum L. as 5 SK 

Heliotropium diversifolium F. Muell. ex Benth a x es ae or 

Heliotropium ethelium Ewart et McLennan oe ae oe a st x 

Heliotropium fasciculatum R.Br. an we a M S< x x 

Heliotropium filaginoides Benth. te Yes Ee x aS ot 

Heliotropium flaviflorum W.V.Fitzg. .. ae ae ae x 

Heliotropium heteranthum (F.Muell.) Ewart et Davies x ae 2 

Heliotropium indicum L. ae = avs a ae x 

Heliotropium ovalifolium Forsk. x x x 
xX x 


90 O6 G5 x 
Heliotropium paniculatum R.Br. ae af an x x 
Heliotropium pleiopterum F. Muell. x 
Heliotropium strigosum Willd. ex DC. .. : X< 
Heliotropium tenurifolium R.Br. var. parviflorum J.M. 

Black .. at x 
Heliotropium tenuifolium R.Br. var. tenuifolium x x 
Heliotropium undulatum Vahl... aa a a Ye aa ae x 
Heliotropium ventricosum R.Br. .. a or At he a x x 
Heliotropium sp. aes a Sy 533 a x at fas ats 
Messersmidia argentea (L.f.) I. M.Johnston .. ate ae ow a 3 
Omphalolappula concava (F. Muell.) Brand. att ok x 
Trichodesma zeylanicum (Burm.f.) R.Br. var. lati- 

sepalum F. Muell. Be ae ie ue iw. Sc x x ig 
Trichodesma zeylanicum (Burm. f.) R.Br. var. zeylanicum x x % x 


ERRETIACEAE 
Ehretia saligna R.Br. ote vs = ae a x »< x 


VERBENACEAE 
Avicennia marina (Forsk.) Vierh. var. resinifera (Forst. f.) 

Bakh. : ae aid te 393 
Callicarpa cana L. oe br ‘a re ots x 
Clerodendrum cunninghamii Benth. an a oh ae hy 
Clerodendrum floribundum R.Br. a i a x x x 
Olerodendrum holtzec F. Muell. 

Olerodendrum tnerme (L.) Gaertn. ae 58 
Clerodendrum tomentosum R.Br. 6 Bu3 bi ifs x 
Denisonia tenuifolia F. Muell. 

Dicrastylis beveridgec F. Muell. 

Dicrastylis costelloi F. M. Bail. 

Dicrastylis dorani F.Muell. var. doranii 

Dicrastylis doranit F.Muell. var. eriantha F. Muell. 

ex J. M. Black ae oe bs 

Dicrastylis ewsuccosa (F.Muell.) Druce .. 

Dicrastylis gilesii F. Muell. ae we 

Dicrastylis lewellini (F. Muell.) F. Muell. 

Gmelina dalrympleana (F. Muell.) H. J. Lam. 

Hualeya linifolia Ewart et Rees 5 
*Lantana camara L. 

Newcastlia bracteosa F. Muell. 

Newcastlia cephalantha F.Muell. .. 

Newcastlia spodiotricha F. Muell. ante ae 
Phyla nodiflora (L.) Greene ys ii a ear ae x 
Pityrodia hemigenioides (F. Muell.). F. Muell. ex Benth. Pe 3h 
Pityrodia jamesii Specht are ag ae oe #8 #- Ms 
Premna acuminata R.Br. .. ale ‘te Ne as ots x Xx 
Premna integrifolia L. 

Premna obtusifoia R.Br. .. ae a oe Sis a. 

Spartothamnella puberulus (F.Muell.) Maid. et Betche Xx 

Spartothamnella teucriflora (F. Muell.) Moldenke 5 ¢ x 
*Stachytarpheta dichotoma (Ruiz. et Pav.) Vahl.. Ei: 
*Stachytarpheta jgamaicensis (L.) Vahl a ; 

Verbena macrostachya F.Muell. .. $3 Bag ae x 

Verbena officinalis L. ie ar oe ax 56 x ap he 
Vitex glabrata R.Br. ys of hi it ae ae ha 4 
Vitex ovata Thunb. . . ; 

Vitex rotundifolia L.f. ca sh Bi te we 36 ae 
Vitex trifolia L. wa ef ay es z. Ye He aye S< 


DOR ORO GEO 2 
5 wNegn OS oXos OS OS EN 


MeN SSS 


— XS SG 
SEIS ee ON OR OS 


xX XXX: 


PROCEEDINGS OF THE LINNEAN SoctetTy or New SourH WateEs, Vou. 96, Part 4 


G. M. CHIPPENDALE 257 


C.A. Baton VebD Dp adaG:. 


LABIATAE 
Anisomeles salviifolia R.Br. ie ais Be Ae AG x 
Basilicum polystachyon (L.) Moench ; A Me 
Coleus scutellarioides (L.) Benth. var. angustifolia Benth. an ¥ x x 
Coleus scutellarioides (L.) Benth. var. laxa Benth. . 

Coleus scutellarioides (L.) Benth. var. limnophila F. Muell. 
ex Benth. : Sai te oe Me iy 
Coleus scutellarioides (L.) Benth. var. scutellarioides 
*Hyptis suaveolens (L.) Poit 
*Leonotis nepetaefolia R.Br. : ane a ae ah 
Mentha australis R.Br... : ae oy Sd x 
Microcorys macredieana F. Muell. ae ae ie x i; a 
Moschosma australe Benth. : : ae LY Ae x 
Ocimum sanctum L. var. angustifolium Benth. ate we sie x 
Plectranthus intraterraneus S8.T. Blake .. eu eA x 
Plectranthus sp. : see me 
Prostanthera baxteri A. Cunn. ex Benth. var. crassifolia 
Benth. ; se a em Bt 
Prostanthera baxteri ie Cunn. ex Benth. var. sericea 
J. M. Black F ae in = x 
Prostanthera striatiflora F. Muell. Be mae os x 
Prostanthera wilkieana F. Muell. M 
Teucrium grandiusculum F. Muell. et Tate x 
Teucrium integrifolium F.Muell. ex Benth. F x 
Teucrium racemosum R.Br. 3% Sih Sg a x 
Wrixonia sp. x 


wa 
“An 


SOLANACEAE 
Anthotroche blackw KF. Muell. : 
Datura leichhardtii F. Muell. ex Benth. 
Duboisia hopwoodi (F. Muell.) F. Muell. 
tLycium ferocissumum Miers aes 
Nicotiana benthamiana Domin 
Nicotiana excelsior J. M. Black 
*Nicotiana glauca Grah. 
Nicotiana gosset Domin 
Nicotiana ingulba J. M. Black : 
Nicotiana megalosiphon Heurck et J. Muell. 55 
Nicotiana occidentalis Wheeler ssp. cg Beers 
Nicotiana simulans Burbidge 
Nicotiana velutina Wheeler = om ae eh aa a 
Physalis minima L.. be ae ae he a. x x 
Solanum asymmetriphyllum Specht. ee hi 28 x 195 
Solanum centrale J. M. Black : Pa a ae 
Solanum chenopodinum F. Muell. a ae see x 
Solanum coactiliferum J. M. Black oe a a uy 
Solanum cunninghamw Benth. .. ie at ae id M: x 
Solanum diversiflorum F.Muell. .. Ae = ok x ee x 
Solanum echinatum R.Br. . . at Le a a Ww x 
Solanum ellipticum R.Br. 
Solanum eremophilum F. Muell. 
Solanum esuriale Lindl. 
Solanum aff. esurtale Lindl. 
Solanum ferocissimum Lindl. 
Solanum lasiophyllum Dun. 
Solanum lucaniw F. Muell. Me ys Ae F ck. oe 
Solanum melanosper mum FF. Muell. a ae a, x SK a x 
Solanum nigrum L. : at ho va x 
Solanum aff. orbiculatum Dun. .. a vs ze Sx os Sf 
Solanum petrophilum F.Muell. var. *. = 2 x ts x 
Solanum phlomoides A. Cunn. ex Benth. on By x6 uit a3 a 
Solanum pugiunculiferum C.T. White .. Wye ts it ke 6 x 
Solanum quadriloculatum F. Muell. oe as ae x x x 
Solanum sturtianum F. Muell. ae & i Se x = S< 
Solanum tetrandrum R.Br... ae vias bat ah ans 
Solanum wilkinsvi S. Moore ae ss sa re Se st es x 


SOSK ES SK SOOKE SC 


Xi 


x > 


ORES OK ORS 6 
x 


ae 


Saar 


PROCEEDINGS OF THE LINNEAN Society or New SourH WaAteEs. Vou. 96, Part 4 


258 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. &G. 


SCROPHULARIACEAE 
Adenosma muellert Benth. 
Bacopa floribunda (R.Br.) Wettst.. 
Buchnera filicaulis O. Schwarz ac iB ae a5 
Buchnera linearis R.Br... e be Hs we x< 
Buchnera tenella R.Br. 
Buchnera tetragona R.Br. 
Buchnera urticifolia R.Br. aA 4 
Centranthera cochinchinensis (Lour.) Merr. A #3 
Elacholoma hornit F. Muell. et Tate ae ae Ss x 
Hemiarrhena plantaginea Benth. .. : 
Tlysanthes clausa (¥. Muell.) With. 
Ilysanthes mitrasacmoides O. Schwarz an ah Sto 
Iimosella australis R.Br. .. oe a ae fe x 
Limnophila chinensis (Osbeck) Merr. at De e: . Ste 
Limnophila fragrans (Forst.f.) Seem. .. we te oie i x 
Lindernia alsinoides R.Br. Me 
Lindernia crustacea (L.) F. Muell. 
Lindernia grossidentata O. Schwarz Are aa ae ae om 
Lindernia lobelioides (F. gece F. Muell. ne ae x ae x 
Lindernia pubescens (Benth.) F. Muell. 28 
Tindernia scapigera R.Br. .. 
Lindernia subulata R.Br. var. glanduligera Specht A oe ea 
Lindernia subulata R.Br. var. subulaia .. a es x 
Microcarpaea muscosa R.Br. 
Mimulus gracilis R.Br... 
Mimulus linearis (R.Br.) Wettst. 
Morgania floribunda Benth. : 
Morgania glabra R.Br. 
Morgania gracilis R.Br. 
Morgania parviflora Benth. 
Morgania pubescens R.Br... 
Peplidium humifusum Delile a 
Peplidium maritimum (L.f.) Wettst. 
Peplidium muellert Benth. 
Scoparia dulcis L. 
Stemodia coerulea Benth. 
Stemodia debilis Benth. ‘ is ke 5a Be He 
Stemodia grossa Benth. .. a8 an Bi is a is x 
Stemodia micrantha O. Schwarz Ae We a ae a oF 
Stemodia viscosa Roxb. .. 3 ae as ee: x ee x 
Stemodia viscosa Roxb. var. x 
Striga curviflora (R.Br.) Benth. : a bh ate oe a 
Striga multifiora Benth. .. ae oy Ae oe x ie x 


M35 2K 5 OX 
xxXXXXXXX 


5 OE 
Dane Sea 


eos OX eX Oe SOE 
x 
x x 
SSS OS ee OOO OOO OGIO 


x 
5 Mo NM G 


XX 5 


BIGNONIACEAE 
Dolichandrone filiformis (DC.) Seem. .. a ai i a x x 
Dolichandrone heterophylla F. Muell. Ae ae i x x< 
Pandorea doratozylon (J. M. Black) J. M. Black a Xx 


PEDALIACEAE 
Josephinia eugeniae F. Muell. ‘e oe ae x he x are 
*Sesamum indicum L. ne ae we a ae ee 6 ae x 


MARTYNIACEAE 
*Martynia annua L... 


x 


LENTIBULARIACEAE 
Utricularia albiflora R.Br.. 
Utricularia bifida L. ex D. Oliver 
Utricularia biloba R.Br. 
Utricularia caerulea L. é 
Utricularia capilliflora F. Muell. 
Utricularia ceratophylloides O. Schwarz x oes Ms ae Pie 
Utricularia chrysantha R.Br. an =o se fea or af ~< 
Utricularia dunstanii Lioyd 


x XX XK XK XK XK XK 


PROCEEDINGS OF THE LINNEAN SootETY oF NEw SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 259 


C.A. Bee 


Utricularia exoleta R.Br. .. ar ah Ss ats ee i ye 
Utricularia flava R.Br. 


Utricularia 


flexuosa Vahl a 


Utricularia fulva F. Muell. 


Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 
Utricularia 


ACANTHACEAE 


hamilton Lloyd 

holtzec F. Muell. : , 
infleca Forsk. var. stellaris (L. 
kamienski F. Muell. 
kimberleyensis C. A. Gardn. 
lasiocaulis F. Muell. 
leptoplectra F. Muell. 
leptorhyncha O. Schwarz 
limosa R.Br. .. 

minutissima Vahl 

pachyceras O. Schwarz 
scandens Benj. 

singerana F. Muell. 

tubulata F. Muell. 

uliginosa Vahl. . 


f.) P. Taylor 


Acanthus tlicifolius L. 
*Andrographis paniculata (Burm.) “Wall. ex Nees 
*Barleria prionitis L. is ae ie F 


Dicliptera glabra Dene. .. 
Dipteracanthus corynothecus ( 


F. Muell. ex Benth.) Brem. 


var. corynothecus : 
Dipteracanthus cor ynothecus (F. Muell. ex Benth. Brem. 

var. grandiflorus Brem. Bio : Ac 
Dipteracanthus sessiliflorus Brem. 


EHbermaiera 
Hygrophila 


glauca Nees 
salicifolia Nees 


Hypoestes floribunda R.Br. var. angustifolia Benth. 
Hypoestes floribunda R.Br. var. floribunda : 
Hypoestes floribunda R.Br. var. paniculata Benth. 
Nelsonia campestris R.Br. oe is 
Rostellularia pogonanthera F. Muell. : 
Sarojusticia kempeana (F. Muell.) Brem. 


Thunbergia 


MYoProRACEAE 


Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 
Hremophila 


arnhemica F. Muell. 


alternifolia R.Br. : 

battii F.Muell. var. battis .. 
battia F. Muell. var. major J. M. Black 
bignoniifiora F. Muell. 
calycina 8. Moore 

castelli arminit H. Pritz. 
christophert F. Muell. 
cordatisepalea L. Smith 
dalyana F. Muell. 

duttont F. Muell. 

eldert F. Muell. 

exotrachys Kraenzl. 


EHremophila freelingii F. Muell. 


Hremophila 
Hremophila 
Hremophila 
Eremophila 
Hremophila 
Hremophila 
EHremophila 
EHremophila 
Hremophila 
EHremophila 


gibsoniit F. Muell. é 
gilestt F. Muell. var. argentea Ewart : 
gilestti F. Muell. filiforme Ewart et Davies 
gilesii F. Muell. var. gilesi .. : L 
glabra (R.Br.) Ostenf. 
goodwintt F. Muell. 

latrobet F. Muell. 
leonhardiana EK. Pritz. 
longifolia (R.Br.) F. Muell. 


macdonnellii EF. Muell. var. glabriuscula 


J. M. Black 


Hremophila 


PROCEEDINGS OF THE LINNEAN SocreTy or NEw SoutH Wares, Vow. 96, Part 4 


PeedanneliD F. Muell. ee, Fananlorornglhs 


5 SOS 5 


SOK OS OK OS OS OS OS OS OS OS CS OS OS OS OS ON OK OS ON ON 


xX X 


Bie ro Grose 0 


x X xxKXKXKKKXKXKXKKKKKKXKXKXKX 


XXXxXXXKX: 


x 6 


x 


260 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D.& G. 


Eremophila macdonnellii F. Muell. var. macrocarpa Ewart 
et Davies ne og als oe: ee 

EHremophila maculata (Ker-Gawl.) F. Muell. 

Eremophila obovata L. Smith var. obovata 

Eremophila paisleyi F. Muell. 

Eremophila rotundifolia F. Muell. ae 

Eremophila serrulata (A. Cunn. ex A. DC.) Druce 

Eremophila strehlowiit HB. Pritz. .. it oH 

Eremophila strongylophylla F. Muell. 

Eremophila sturtu R.Br. .. 

Eremophila willsiia F. Muell. var. integrifolia Ewart 

Eremophila willsit F. Muell. var. willsi 

Myoporum montanum R.Br. 


SK oN 2S DE ORK OK OOK OK ON KX 


PLANTAGINACEAE 
Plantago varia R.Br. sens. lat. 


MZ 
A 


RUBIACEAE 
Borreria australiana Specht ‘ ae hs x x 
Borreria brachystema (R.Br. ex Benth.) ‘Valet... Ae x x 
Borreria breviflora (F. Muell. ex Benth.) Specht ab ce x 
Borreria breviflora (F. Muell. ex Benth.) Specht var. si ch 
Borreria carpentariae Specht as “igs £; 
Borreria euserta (Benth.) K. Schum. 5 ae 
Borreria geeingwa Specht var. dilatata (Benth.) “Specht 
Borreria geeingwa Specht var. gaa em ) mp 
Borreria gilliesae Specht : a oe 
Borreria hillit Chippendale ea x x ats be 
Borreria tnaperta (F. Muell.) K. Schum.. : 5 ae Shs < 
Borreria involucrata (O. Schwarz) Chippendale 2 
Borreria leptoloba (Benth.) Specht 5 iF ok Ae a: a 
Borreria pogostoma (Benth.) K.Schum... ae ee of ao x 
Borreria suffruticosa Spey fe 2 Si 2 a 7 
Borreria sp. ae age oe ae x x 
Canthium attenuatum R.Br. ex Benth. 
Canthium latifolium F. Muell. ex Benth. 
Canthium lineare E. Pritz. oe an Bs sft se: be: 
Canthium aff. lucidum Hook. et Arn. “2 tt Le Me o): ae x 
Canthium schultzii (O. Schwarz) Chippendale .. i te a i x 
Canthium sp. nie as = x ‘ve a 
Coelospermum reticulatum (F. Muell. ) Benth. .. ee ne sad x x 
Dentella asperata Airy-Shaw ae ie ah oe x ie it bs 
Dentella browniana Domin oe : ms Ee BY bs uff x 
Dentella dioeca Airy-Shaw te si = x 
Dentella pulvinata Airy-Shaw var. “repanda Airy- Shaw x in vi 
Dentella repens (L.) Forst. et f. 2 sh fy x x 
Gardenia edulis F.Muell. .. : mE ae ae ie os SE x 
Gardenia fucata R.Br. ex Benth. x 
Gardenia megasperma F.Muell. var. arborea Ewart et 
Cookson Bs ina x x 
Gardenia megasperma “F. Muell. var. megasperma a: fe oe x x 
Gardenia aff. megasperma F. Muell. ae i He fi Ee Shs x 
x 
x 


x X X 


XXX KKK KK XK 


KR OX 


x X X 


Gardenia petiolata O. Schwarz ves a6 sity a AY 
Gardenia pyriformis A.Cunn. ex Benth. ai ta a as x 
Gardenia resinosa F. Muell. : ae es bs a oe x 
Gardenia suffruticosa R.Br. ex Benth. .. veh ae ae Ste a x 
Guettarda speciosa L. fs of Sid son we a 
Hedyotis mitrasacmoides F. “Muell. oo Je ae ah un < x 
Hedyotis pterospora F. Muell. aie alg ae be 4 oe 4 
Hedyotis scleranthoides F.Muell. .. a8 burt a x ae x 
Ixora coccinea L.  . ae 

Ixora klanderana F. Muell. 

Izxora pentamera Benth. 

Izora timorensis Dene. gy as oe Ae eke Bis ui 
Ixora tomentosa Roxb. sts a a ae a ah ie x 
Knouia stricta Gaertn. 


OKO 


PROCEEDINGS OF THE LINNEAN SocrETY or NEw SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 261 


C.A. B.T. V.R.D. D. &G. 


Morinda citrifolia L. in ae Oe Ae 2 ie Bc 
Nauclea orientalis L. se ae we zie Bn a8 st x 
Oldenlandia corymbosa L. .. ae to sis # a 

Oldenlandia galioides (F. Muell.) F. Muell. Sg a x 

Oldenlandia mollugoides O. Schwarz ae : 

Oldenlandia tenuissima O. Schwarz 

Pavetia brownw Brem. var. brownti 

Pavetta browni Brem. var. glabra Brem. a Be ae of 
Pavetta granitica F.Muell. ex Brem. .. ae ys or x x 
Pavetta muellert Brem. . vy fu 

Pomaz umbellata (Gaertn.) Soland. ex A. ‘Rich. ay x 

Psychotria nesophila F. Muell. ; : ie 

Randia cochinchinensis (Lour.) Merr. 

Randia lamprophylla O. Schwarz 

Scyphiphora hydrophyllacea Gaertn. f. .. Sa ve she 

Synaptantha tillaeacea (F. Muell.) Hook. f. a ae x 

Tarenna dallachiana (F. Muell. ex Benth.) 8S. Moore .. an a 35 
Timonius timon (Spreng.) Merr. .. ss a Ba aa ae x 


XXX XK XX XK X 


> 3 


SS KE 


Sos 


CUCURBITACEAE 
Bryonopsis laciniosa (L.) Naud. .. 
*Citrullis colocynthus (L.) Schrad. 
*Citrullis lanatus ae Mansf. 
Coccinea cordifolia (L.) Cogn. 
Coldenia procumbens L. 

Cucumis melo Li. ssp. agrestis (Naud.) Greb. 
*Cucumis myriocarpus Naud. : my 8 Se nt 
Luffa aegyptiaca Mill. ae Sy me ke 3 ae te oe x 
Luffa graveolens Roxb.  .. oo Be sf 0 ae x 90 
Melothria maderaspatana (L.) Cogn. : Ae x x x x 
Melothria micrantha (F. Muell.) F _ Muell. ex Cogn. Bs x : 

Trichosanthes cucumerina L. : De zh i x 


xe 
x 
x 


ONION 


xX: 


CAMPANULACEAE 

Isotoma petraea F.Muell. .. a) RS ag aS x 4 Pe 

Lobelia arnhemiaca Wimmer a ye ae fs ae a8 x ate 
Lobelia dioica R.Br. 3 ae oi = sis ae bs O60 x 
Lobelia heterophylla Labill.. ‘ ae Re a ae ¥ ote 
Lobelia stenophylla Benth. a a a ak = a ap x 
Lobelia quadrangularis R.Br. a oe aa ae % a) XK x 
Lobelia aff. Migr yokes de Vr. 
Wahlenbergia sp. 
Wahlenbergia sp. 
Wahlenbergia sp. 
Wahlenbergia sp. .. ae a Be a ae A 
Wahlenbergia sp. .. Pe 2 sts a ae oh oo x Ss 
Wahlenbergia sp... a4 au Mi 5 ae iM we No x 


x 


XXX X X 


GOODENIACEAE 
Calogyne berardiana (Gaud.) F.Muell. .. mite ae x 
Calogyne heteroptera F. Muell. : Be Be ras a 
Calogyne hians O. Schwarz 
Calogyne holizeana Specht is us ae Pe » 
Calogyne pilosa R.Br. ie as ae oe te of x 
Calogyne purpurea F. Muell. me oe Ay et oN 
Calogyne raphanoides O. Schwarz. . a 
Catosperma goodeniaceum (F'. Muell.) Krause 
Dampiera candicans F. Muell. 
Dampiera cinerea Ewart et Davies 
Goodenia armitiana F.Muell. var. armitiana .. : 
Goodenia armitiana EF. Muell. var. multicaulis Blakely : 
Goodenia armstrongiana de Vr. .. ae su ay wis oe x x 
Goodenia aff. armstrongiana de Vr. ie ae fs ae ae ok SS 
Goodenia auriculata Benth. 5 oe AS ws 22 = x x 
Goodenia aff. auriculata Benth. .. an i is ae oe * x 
Goodenia azurea F. Muell. .. as ne ue a x x S< 


Be ON Ew OK OK KOK 


XK KKK: 


PROCEEDINGS OF THE LINNEAN Society or New SoutrH Wates, Vou. 96, Part 4 


262 


NORTHERN TERRITORY PLANTS 


Goodenia 
Goodenia 
Goodenta 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 


basedowtt Krause 

cirrifera F. Muell. ays 

aff. coronopifolia R.Br. 

cycloptera R.Br. .. nie 

erecta Hwart 

glabra R.Br. we 

glauca F. Muell. .. 

gracilis R.Br. 

grandiflora Sims 

heterochila F. Muell. var. ‘foliosa Benth. 
heterochila F. Muell. var. heterochila .. 
heterochila F. Muell. var. runcinata Benth. 
hirsuta F. Muell. 

hispida R.Br. ; 

aff. hispida R.Br. 

horniana Tate .. 

lamprosperma F. Muell. 

larapinta Tate .. 

linifolia W. V. Fitzg. ex ; Krause 
lunata J. M. Black : 

microptera KF. Muell. 

mitchelliana Benth. 

mueckeana F. Muell. 

pumilio R.Br. 


purpurascens R.Br. var. minima F. Muell. ex 


Benth. 


Goodenia 
Goodenia 
Goodenia 


Goodenia sepalosa F. Muell. ex Benth. var. brachypoda 


purpurascens R. Br. var, “purpurascens 
ramelii F. Muell. : 
scaevelina F. Muell. 


F. Muell. ex Benth. 


Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 
Goodenia 


sepalosa F. Muell. ex Benth. var, sepalosa Bh 


strangfordi F. Muell. 
subintegra F. Muell. ex Tate 
vilmorinae F. Muell. 

sp. 

s 


Pp: 
Leschenaultia agrostophylla. F. Muell. 
Leschenaultia divaricata F. Muell. 
Leschenaultia filiformis R.Br. 
Leschenaultia striata F. Muell. 


Scaevola 
Scaevola 
Scaevola 
Scaevola 
Scaevola 
Scaevola 
Scaevola 


aemula F. Muell. .. 
aff. aemula F. Muell. 
angulata R.Br. .. 
collaris F. Muell. .. 
daleana Blakely .. 
densevestita Domin 
depauperata R.Br. 


Scaevola frutescens (Mill.) Krause 


Scaevola 
Scaevola 
Scaevola 


ovalifolia R.Br. var. glabra R. Br. 
ovalifolia R.Br. var. ovalifolia 
paniculata Ewart et Davies 


Scaevola parvifolia F. Muell. ex Benth. 


Scaevola 
Scaevola 


revoluta R.Br. 
spinescens R.Br. 


Velleia connata F. Muell. 
Velleta glabrata Carolin 
Velleia macrocalya% de Vr. 


BRUNONIACEAE 
Brunonia australis Sm. 


STYLIDIACEAE 
Levenhookia chippendalet Erickson et Willis 
Stylidium alsinoides R.Br.. 
Stylidium ceratophorum O. Schwarz 


C.A. 


DX 


XS OK 06K 


x 6 


a XX DS SS Kee 


xXx xX: 


x 


B.T. 


4 OX KOE 


V.R.D. D. & G. 


o¢ x 
x O¢ 
x 
x oe 
ac x 
x 
x 
x 
x x 
x 
x 
x x 
x x 
x 
x oc 
6 0 x 
x 6.0 
x 
x x 
x 
x 
x 
x x 
x ono 
26 x 
x o¢ 
x 
x 
x 
Xx xX 
O06 x 


PROCEEDINGS OF THE LINNEAN SocrETY oF New SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 263 


C.A. 1eyib | (WV ARID, ID). ay (GC 


Stylidium cordifolium W. V. Fitzg. 
Stylidium ericksonae J. H. Willis .. i wt BE Fa! ae ic 
Stylidium fissilobium F.Muell.  .. oa re Sil * e x 
Stylidium floodi F. Muell. ye me 

Stylidium floribundum R.Br. 
Stylidium imaequipetalum J. M. Black 
Stylidium irriguum W. V. Fitzg. Bee a8 ne ae a 
Stylidium leptorrhizum F. Muell. des te om on se x 
Stylidium lobuliflorum F. Muell. .. Be “a as oe & x 
Stylidium multiscapum O. Schwarz ot ot a at se 
Stylidium muscicola F. Muell. a We fi a 2G Me 4 
Stylidium pachyrrhizum F. Muell. 

Stylidium pedunculatum R.Br... 
Stylidium quadrifurcatum Hrickson et Willis 
Stylidium schizanthum F. Muell. 

Stylidium tenerrimum F. Muell. 

Stylidium uliginosum Sw. 


5 ON OX OX 
< 
XX XK XK XK 


XxX! 


eee 
ZNvie 


XxX XXXXXX: 


CoMPOSITAE 

*Acanthospermum hispidum DC. .. =p ve te: an 

Actinobole uliginosum (A. Gray) Hichler. . 3 BS x 

Adenostemma lavenia (L.) Kuntze ; ‘ 

Ageratum conyzoides L.  . ae 

Angianthus pusillus (Benth.) Benth. 

Angianthus tenellus (F. Muell.) Benth. 

Bidens bipinnata L. * 53 

*Bidens pilosa L. .. 

Blainvillea dubia Specht . : 

Blumea diffusa R.Br. ex Benth. 

Blumea hieracifolia (D. Don) Dey ss Ae & ys 

Blumea integrifolia DC. .. ii He £6 lf ts x 

Blumea lacera (Burm. f.) DC. 

Blumea mollis (D.Don) Merr. 

Blumea pannosa O. Schwarz 

Blumea sazatilis Zoll. et Mor. 

Brachycome blackiit Davis : 

Brachycome ciliaris (Labill.) Less. var. ciliaris. 

Brachycome ciliaris (Labill.) Less. var. lanuginosa (Steetz) 
Benth. .. ae ae af ae 

Brachycome iberidifolia Benth. 

Brachycome lineariloba (DC.) Druce 

Brachycome tesquorum J.M. Black 

Calocephalus knappii Ewart et Davies . 

Calocephalus platycephalus ee aca Benth. = oF . 

Calotis breviseta Benth.  . as cud S x x x 


5s eX 
OS 6 OS 


x X 5 MM 
5 OS OS OS Gg 
5 ON OS ON ON ON ON 0% ON OS OS G 


xX XK XK XK & 


Calotis cuneifoa R.Br... x 

Calotis cymbacantha F. Muell. x 

Calotis erinacea Steetz os on x i 

Calotis hispidula (F. Muell.) F. Muell. x x 4 
Calotis kempei F.Muell. .. x 

Calotis latiuscula F. Muell. et Tate x 

Calotis multicaulis (Turez.) Druce X< 

Calotis porphyroglossa F. Muell. ex Benth. x 
*Carthamus lanatus L. a x A: 
Centipeda cunninghamiu (DC.) A.Br. et Aschers x Xx 
Centipeda minima (L.) A.Br. et Aschers var. lanuginosa 

(DC.) Domin ae 3 os: ae eh x = 

Centipeda minima (L.) A.Br. et Aschers var. minima x S 
Centipeda thespidioides F. Muell. x 

Chthonocephalus pseudevax Steetz x as 
Coleocoma centaurea F. Muell. x x 
*Conyza bonariensis (lu.) Cronquist ‘ Lit Sis x oe rs 
Craspedia chrysantha (Schlechtd.) Benth. a on ck x 

}+Cryptostemma calendula (l.) Druce 3 ae SS a8 

Eclipta alatocarpa Melville ; te ae ae a S bu St 
Eclipta platyglossa F. Muell. a gs a a si es ae x 


PROCEEDINGS OF THE LINNEAN Society or New SoutrH WaAtzEs, Vou. 96, Part 4 


264 NORTHERN TERRITORY PLANTS 


C.A. B.T. V.R.D. D. &G. 


Eclipta prostrata (l.) L. 
Hlephantopus scaber Bx : 
*Hmilia sonchifolia (L.) DC. os ys 5 ae 
Epaltes australis Less. : a G6 be he 4 Le x 
Hrigeron ambiguus F. Muell. ee 45 nea ne a x 
*Hrigeron floribundus (H.B.K.) oe ae 
Flaveria australasica Hook. on bcs 
Glossogyne filifolia F. Muell. ex Benth. oa a a we x 
Glossogyne tenuifolia (Labill.) Cass. ie mae ie a 
Gnaphalium involucratum Forst. f. 

Gnaphalium luteoalbum L. ae 

Gnephosis eriocarpa (F. Muell.) Benth. 

Gnephosis foliata (Sond.) Hichler ae 

Helichrysum ambiguum Turez. var. ambiguum . 
Helichrysum ambiguum Turcez. var. paucisetum J. M. Black 
Helichrysum aff. ambiguum Turcz. 


a OS 2X OX OS OX 


xX X 
x 
x 5 
x 


Dom 


Helichrysum 
Helichrysum 
Helichrysum 
Helichrysum 
Helichrysum 
Helichrysum 
Helichrysum 
Helichrysum 
Helichrysum 


apiculatum (Labill.) D 
ayersit F. Muell. 
bracteatum (Vent.) Andr. 
cassinianum Gaud. . 
davenportit F. Muell. 
kempet F. Muell. 
ramossisimum Hook. 
semifertile F. Muell. .. 
thomsoniit F. Muell. .. 


Helipterum charsleyae F. Muell. 
Helipterum corymbiflorum Schlechtd. 
Helipterum fitzgibbonit F. Muell. .. 
Helipterum floribundum DC. 


Helipterum 
Helipterum 
Helipterum 
Helipterum 
Helipterum 
Helipterum 


moschatum (A. Cunn. ex DC.) Benth. 
pterochaetum (EF. Muell.) Benth. 

saxatile Wilson 

stipitatum (F. Muell.) ¥. Muell. ex Benth. 
strictum (Lindl.) Benth. : aa¢ 
tietkenswi F. Muell. 


tHypochoeris glabra L. : 
Ixiolaena leptolepis (DC.) Benth. : 
¢Lactuca serriola L. var. integrifolia Gren. et Godr. 


ftLactuca serriola L. var. serriola .. 
Millotia greevesw 


F. Muell. ssp. kempet (F. “Muell.) 


Schodde var. helmsii (F. Muell. et Tate) Schodde 


Millotia greevesti 
Schodde var. kempew 


F. Muell. ssp. kempez (F. Muell.) 


Minuria cunninghamti Benth. 

Minuria denticulata (DC.) Benth. 
Minuria integerrima (DC.) Benth. 
Minuria leptophylla DC. .. a 
Moonia ecliptoides (F. Muell.) Benth. 
Moonia procumbens (DC.) Benth. 
Myriocephalus rudallai (F. Muell.) Benth. 


Myriocephalus stuartii (F. Muell. 


Benth. 


et Sond. ex Sond.) 


Olearia ferresii (F. Muell.) ¥. Muell. ex Benth. 
Olearia aff. muellert (Sond.) Benth. : 
Olearia stuartii (F. Muell.) F. Muell. ex Benth. 
Olearia subspicata (Hook.) Benth. : Bi: 
Phacellothrix cladochaeta F. Muell. 

Pluchea dentex R.Br. ex Benth. 

Pluchea indica Less. a le a ie 
Pluchea rubelliflora (F. Muell.) B. L. Robinson 
Pluchea squarrosa Benth. : ae 
Pluchea tetranthera F. Muell. var. tetranthera 
Pluchea tetranthera F. Muell. var. tomentosa Benth. 
Pleurocarpaea denticulata Benth. . . 

Podocoma cuneifolia R.Br... 

Podocoma sp. 


KKK KKKKKKKKKKKKKKKKKKKKKKXXKKXKXKXXX 


x 


0 ON KKK 


xx: 


PROCEEDINGS OF THE LINNEAN SocreTy or New Soura WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 265 


C.A. Bl VER D2 D&G: 


Podolepis canescens A. Cunn. ex DC. 
Podolepis capillaris (Steetz) Diels 
Podolepis georgei Diels ie 
Pterigeron adscendens Benth. a 
Pterigeron cylindriceps J. M. Black 
Pterigeron decurrens (DC.) Benth. 
Pterigeron dentatifolius F. Muell. 
Pterigeron liatroides (Turcz.) Benth. : ye S. 
Pterigeron odorus (F. Muell.) Benth. var. “major ‘Benth. : ats ns x 
Pterigeron odorus (F. Muell.) Benth. var. odorus ; ; 
Pterocaulon glandulosum (F.Muell. ex Benth.) Benth. 
et Hook. f. var. glandulosum a 
Pterocaulon glandulosum (F.Muell. ex Benth.) ‘Benth. 
et Hook. f. var. velutinum Ewart et Davies Se x 
Pterocaulon serratum O. Schwarz .. A ae 
Pterocaulon sphacelatum (Labill.) Benth. et Hook. fie x 
Pierocaulon verbascifolium F. Muell. Bd ae 
Rutidosis helichrysoides DC. i Ee a ae x x 
Senecio aff. cunninghamii DC. 
Senecio gregortt KF. Muell. ae at ae 
Senecio laceratus (F. Muell.) Belcher nt x 
Senecio lautus Forst. f. ex Willd. aff. =P, dissectifolins 
Ali F ie : a3 ae x 
Senecio magnificus F. Muell. tes Be a ve x 
Senecio odoratus Hornemann ae Be oe x ss 
Senecio sp. .. a wits se oe ne x 
Sigesbeckia microcephala DC. a5 iS ie ie x 
*Sonchus oleraceus L. ik Fa ee a x 
Sphaeranthus africanus 188 30 as ae: its Ke oe an: 
Sphaeranthus hirtus Willd. ee sis 5 a ee oe x 
Sphaeranthus indicus L. .. a! 3 wh a By x x 
Spilanthes grandiflora Turcz. 
Synedrella nodiflora (L.) Gaertn. ah 
Thespidium basiflorum (F. eg tee F. Mane 
*Tridax procumbens _L. : ne Et a oe 3 
Vernonia cinerea (L.) Less. var. cinerea ber ee i x >< 
Vernonia cinerea (L.) Less. var. lanata Kost. .. oF x ay ws 
Vittadinia brachycomoides F. Muell. Bs si - a ae x 
Vittadinia macrorhiza (DC.) A. Gray... 3 ie Me x 
Vittadinia pterochaeta (F. Muell. ex Benth.) J. M. Black 
Vittadinia scabra DC. ’ : =e ae 
Vittadinia aff. scabra sensu J. M. “Black 
Vittadinia triloba (Gaud.) DC. sens. lat. 
Vittadinia sp. ae ate : 
Vittadima sp. Boe 
Waitzia acuminata Steetz .. 
Waitzia citrina (Benth.) Steetz 
Wedelia asperrima (Dene.) Benth. : Ne ake ash 
Wedelia biflora (L.) DC. .. te ce he a ne aN ats x 
Wedelia stirlingi Tate a wi ae oe Bc at So Cr 
Wedelia verbesinoides F. Muell. ex Benth. on oe ine re at x 
*Xanthium spinosum L. ae ah oe : ae x 


x XXX XX XX 


> a 
N 
Wa 


ww 
aA 
>. 


exe 


XI: 


SK 


a SOK GX NGS 


xxx KK XK XXX 


x 


ADDENDUM 


Since this paper was submitted to the Society, several papers concerning Northern Territory 
plants have been published. These are: 


Court, A. B., 1971—A new combination in the genus Bossiaea Vent. (Papilionaceae). 
Muelleria, 2 (2): 139-42. 


FRYXELL, P. A., and Hasumt, SHARIFUL H., 1971.—The segregation of Radyera from Hibiscus 
(Malvaceae). Bot. Gaz., 132 (1): 57-62. 


Maconocuigz, J. R., and Byrnes, N., 1971.—Additions to the Flora of the Northern 
Territory. Muelleria, 2 (2): 133-37. 


Maconocuig, J. R., and Parker, 8S. A., 1971.—Further collections of two little known 
Stylidiaceae from the Northern Territory. Muelleria, 2 (2): 145. 


PROCEEDINGS OF THE LINNEAN Socrrty oF NEw SoutH WaAtLEs, Vow. 96, Part 4 


266 


The information and records in these papers will mean small corrections to Tables 1 and 2, 
but do not significantly affect the comments on these tables. 


NORTHERN TERRITORY PLANTS 


total numbers of plant species now known to the following : 


Northern Territory : 


2736 native spp., 114 introduced spp. 


The records in the papers bring the 


Central Australia : 1208 5 on 55 53 
Barkly Tableland : 495 ,, a 8 a 
Victoria River District: 919  ,, 53 9 06 
Darwin and Gulf: 1581 3 AD 64 35 


Additions to the Check List are: 


TYPHACEAE 
Typha orientalis Presl 


HYDROCHARITACEAE 
Blyxa auberti Rich. a 
Maidenia rubra (W. V. Fitzg.) Rendle 


GRAMINEAE 
Eragrostis australasica (Steud.) C. E. Hubbard 
Hragrostis clelandw 8.'T. Blake Ste : 
Sporobolus elongatus R.Br. 


PONTEDERIACEAE 
Monochoria hastata (L.) Solms. 


ORCHIDACEAE 
Nervilia discolor (Bl.) Schltr. ae 
Nervilia holochila (F. Muell.) Schltr. 


PROTEACEAE 
Grevillea erythroclada W. V. Fitzg. 
Grevillea pterosperma F. Muell. 
Hakea rhombales F. Muell.. . 


SANTALACEAE 
Santalum album L. 


CHENOPODIACEAE 
Atriplex suberecta I. C. Verdoorn 
Bassia astrocarpa F. Muell. 
Bassia articulata J. M. Black oi ad oe 
Bassia brachyptera (F. Muell.) R. H. Anderson 
Bassia george: EK. H. Ising ; alps : 
Bassia minuta E. H. Ising 
Kochia radiata P. G. Wilson : ae i 
Malacocera tricornis (Benth.) R. H. Anderson 
Suaeda australis (R.Br.) Mogq. 


AMARANTHACEAE 
Ptilotus royceanus Benl 


PORTULACACEAE 
Anacampseros australiana J. M. Black 
Calandrinia disperma J.M. Black 


MIMOSACEAE 
Acacia pachyacra Maiden et Blakely 


PAPILIONACEAE 


Bossiaea bossiaeoides (A. Cunn. ex Benth.) A. B. Court 


(syn. Bossiaea phylloclada F. Muell.) 
Crotalaria alata Buch.-Ham. ex D. Don 
Crotalaria quinquifolia L. 


ZYGOPHYLLACEAE 
Tribulus hirsutus Benth. 


V.R.D. D. & G. 


x X X 


x XX XK xX Xx 


pa oe 


xX X 


x 


PROCEEDINGS OF THE LINNEAN SocreTy oF NEw SoutH WALES, Vou. 96, Part 4 


G. M. CHIPPENDALE 267 


C.A. Jejdlo WeliID), IDE a's (Cy 


RUTACEAE 
Geijera linearifolia (DC.) J. M. Black 


x 


EUPHORBIACEAE 
Omalanthus populifolius Grah. .. sets 6 vis ait ee oo Ke 


RHAMNACEAE 
Eimmenosperma cunninghami Benth.  .. 56 ore ae ae Be x 


MALVACEAE 
Radyera farraget (F. Muell.) Tete et Hashmi ee 
Hibiscus farraget F.Muell.) . x 


CoMBRETACEAE 
Terminalia crassifolia Exell : ae ite are oye a ays x 
Terminalia fitzgeraldi C. A. Gardner 2% whe ae af see ot x 


BoRAGINACEAE 
*Amsinckia hispida (Ruiz. et Pav.) I.M.Johnston .. x 


VERBENACEAE 
Newcastlia cladotricha F.Muell. .. ae ae fe x 


SCROPHULARIACEAE 
Mimulus prostratus Benth. jie sis be ae x 
Mimulus repens R.Br. fe st a oe ae x 


MYOPORACEAE 
Hremophila polyclada (F. Muell.) F. Muell. S33 4 x 
Hremophila turtoni F. Muell. fe re ap Se x 


GooDENIACEAE 
Leschenaultia helmsii Krause a ae “ a x 


STYLIDIACEAE 
Levenhookia chippendalei Hrickson et Willis se “ies x ae 4 ae 


PROCEEDINGS OF THE LINNEAN Soorety or NEw SoutH Watgs, Vou. 96, Part 4 


A BRACHYOPID LABYRINTHODONT FROM THE LOWER TRIAS 
OF QUEENSLAND 


ANNE HowIe 


Tinnean Macleay Fellow 
School of Biological Sciences, University of Sydney* 


[Accepted for publication 21st July 1971] 
(Plates XIV, XV) 


Synopsis 
The skull of a new species of brachyopid labyrinthodont from the Lower Triassic Rewan 
Formation of Queensland is described and compared with other brachyopids using the method of 
Welles and Estes (1969). It is the most complete and the best ossified brachyopid yet found 
and is closely related to a form from the Mangali Beds of Central India. 


INTRODUCTION 


Following an initial productive field trip to the Lower Triassic Rewan 
Formation in 1969 (Bartholomai and Howie, 1970; Howie, in press), a second 
trip was organized in June, 1970. The purpose of this was first to collect from 
known vertebrate-bearing localities, and second to explore likely areas of the 
Bowen Basin for further Rewan exposures. The trip was a success on both counts, 
yielding an excellent capitosaur skull (an account of which is almost ready for 
publication), several small reptile skulls similar to those reported by Bartholomai 
and Howie (1970), a tiny (2 cm.) temnospondylous labyrinthodont skull, and 
many fragmentary fish, amphibian and reptilian remains from the chief known 
locality (Queensland Museum field locality L78), and providing us with a new 
productive locality some 79 miles north-north-east from L78. 


Field Locality 


The new locality (A. A. Howie Field Locality Q6) is at the headwaters of 
Duckworth Creek, south-west of the settlement of Bluff, and lies on the north-west 
edge of the Mimosa Syncline. Here faces of the rarely exposed Rewan Formation 
are found beneath the more massive escarpments of the Middle Triassic Clematis 
Sandstone and the overlying Lower Jurassic Precipice Sandstone which form the 
northern edges of the Blackdown Tableland. Malone, Olgers and Kirkegaard 
(1969) note that in this area the Rewan sediments are so similar lithologically 
and in stratigraphic position to the Rewan Formation of the type area (which is 
near locality L78) that direct correlation is justified and add that they are almost 
certainly continuous below the surface. 


This Duckworth Creek exposure consists of several small linked erosion 
gullies, and specimens are found in these and in the alluvium at their bases. 
Rewan mudstone was exposed to a maximum depth of about 20 feet and included 
a Single broad pale green band of very fine-grained sandstone. 


Associated Material 


Associated with the brachyopid labyrinthodont described below were 
several other amphibian species but no fish or reptiles, a strong contrast with 
locality L78, where reptiles were a common component of the fauna. Laby- 


* Present address, School of Geology, University of Melbourne. 


PROCEEDINGS OF THE LINNEAN Society oF New South WaAtzEs, VoL. 96, Part 4 


ANNE HOWIE 269 


rinthodont remains include a femur which probably belonged to a very large 
capitosaur, left and right femora and other postcranial remains of a more 
terrestrial type of labyrinthodont, nodules containing rhachitomous vertebrae, 
bone scraps bearing ornament of the type described by Cosgriff (1965) in 
Deliasaurus kimberleyensis, numerous coprolites, and fragments of an unprepared 
skull which I think can be assigned to Rewana sp. (Howie, in press). Although 
the brachyopid skull was found in the red mudstone, all the other more complete 
remains came preserved in nodules from the pale green band. 


The Family Brachyopidae 

In 1956 Watson published a review of the brachyopid labyrinthodonts 
and was followed by Cosgriff (1969), who largely used Watson’s criteria for 
inclusion of a species within the Family Brachyopidae, and Welles and Estes 
(1969), who built on studies of Watson, Shishkin (1964, 1966) and Bystrow 
(1935) to establish a revised and extended set of brachyopid characters. 


Within the ‘ brachyopids proper’? Watson recognized Bothriceps australis 
and B. major (which he renamed Trucheosaurus), Brachyops laticeps, Batracho- 
suchus watson and B. brown, Pelorocephalus, and “ Platyceps’’ wilkinsoni, 
but he included forms such as Hobrachyops, Dvinesaurus and the plagiosaurs 
within a larger brachyopid grouping. Panchen showed in 1959 that the 
plagiosaurs were not closely allied to the brachyopids. Cosgriff made ‘“‘ Platyceps”’ 
wilkinsoni the type of a new genus, Blinasaurus which had two known species, 
B. wilkinsoni and B. henwoodi, and recognized Bothriceps, Trucheosaurus, 
Brachyops laticeps, Batrachosuchus watsom, B. browni, Hadrokkosaurus and 
Boreosaurus aS being true members of the Family Brachyopidae. Welles and 
Estes took a more conservative viewpoint, eliminated genera which they thought 
were doubtful (Boreosaurus, for example) and ended with Blinasaurus henwoodi, 
Bothriceps (which included B. australis, Watson’s Trucheosaurus major and 
‘“* Platyceps”’ wilkinsoni, Cosgriff’s type for Blinasaurus), Batrachosuchus watson, 
B. browni and B. lacer (from Sushkin’s Batrachosuchoides lacer), Brachyops and 
Hadrokkosaurus, thus reducing the number of brachyopid genera to five. 


I intend to follow Welles and Estes’ grouping of the brachyopids and their 
set of brachyopid skull characters, while adding to these Cosgriff’s characters 
for brachyopid lower jaws. I do not agree with Cosgriff’s brachyopid skull 
character of palatal teeth being present only on the vomer bones; irregular 
palatine teeth can be seen in Hadrokkosaurus (on the ectopterygoid and palatine 
bones) and in Brachyops allos nu. sp. (on the ectopterygoid bone). 


SYSTEMATIC DESCRIPTION 

Class Amphibia 
Subclass Labyrinthodontia 
Order Temnospondyli 
Superfamily Brachyopoidea 
Family Brachyopidae 

Characters of the family. Skull short, broad ; no zones of intensive growth 
(other than in the cheek region—see below). Orbits anterior and usually 
relatively large. Otic notch absent, or at most a shallow embayment. Tabulars 
short and broad; tabular horns absent or weak. Parasphenoid flat, becoming 
elevated anteriorly above vomers. Usually a single tusk pit pair on vomer, 
palatine, and ectopterygoid ; interstitial smaller teeth little developed or absent ; 
tusks usually much elongated and massive ; dentary tooth row relatively short. 
Vomerine plate short. Occiput with strong slope posteroventrally to occipital 
condyles, the latter usually quite large. Quadrate condyles large, ventrally 
produced, anteroventral in position relative to occipital condyles. Squamosal 
and qaadratojugal with strong occipital flanges forming a vertical transversely 


PROCEEDINGS OF THE LINNEAN Socipty oF New SoutH Wages, Vou. 96, Part 4 


270 A BRACHYOPID LABYRINTHODONT 


concave trough lateral to the pterygoid. Pterygoid forming a steeply arched, 
flat-roofed palate. Quadrate somewhat compressed laterally, wedged antero- 
laterally between flange of pterygoid and squamosal-quadratojugal trough. 
Retroarticular process elongate. Posterior meckelian foramen and angular- 
prearticular suture on ventral surface or very low on lingual surface. 


Brachyops Owen 1855 


Type species: Brachyops laticeps Owen. Brachyops is the first described 
and thus the typical brachyopid. Other brachyopids differ from Brachyops 
mainly in their skull proportions—the most obvious differences are as follows : 


Blinasaurus Cosgriff (1969) differs in that its snout is more rounded, its 
orbits are larger and therefore closer together, its external nostrils are closer 
together, its parietal foramen is relatively much closer to its orbits, its tabulars 
and postparietals are reduced, its interpterygoid vacuities are much longer 
relative to their width, so that the anterior part of the palate is shortened and the 
posterior part lengthened. 


Bothriceps Huxley (1859) has a much narrower skull so that its orbits and 
nares are closer together, and its interpterygoid vacuities are longer relative to 
their width. 


Hadrokkosaurus Welles (1957) is larger than Brachyops, its skull is much 
broader relative to its length and is shorter postorbitally. 


Batrachosuchus Broom (1903) has a broader, less pointed skull, the preorbital 
skull is shorter, the interpterygoid vacuities are longer relative to their width, 
and the anterior part of the palate is shortened. 


Brachyops allos n.sp. 


Holotype. A skull complete but for the left cheek region, part of the right 
quadratojugal and pterygoid, and a few scraps from the dermal skull roof. 
Queensland Museum No. F6572. 


Type locality. A. A. Howie field locality Q6, near the headwaters of the 
Duckworth Creek, south-west of the settlement of Bluff, South Central 
Queensland. 


Horizon. Lower Upper Rewan Formation of the Mimosa Group, Lower 
Trias. 

Characters of the species. Brachyops allos differs from Brachyops laticeps, 
especially in that its exoccipital condyles are much nearer the level of the quadrate 
condyles, so that the backwardly sloping portion of the occiput in B. allos is 
greatly reduced. Although this character is one which Watson (1956) considered 
showed the stage of evolutionary advancement of a brachyopid, the two species 
are otherwise so alike that generic separation at this stage would be foolish. The 
cultriform process of the parasphenoid in B. allos is clasped laterally by posteriorly 
directed processes of the vomers rather than overlying these as it does in B. 
laticeps. Posteromedially the process bears an area of dermal denticles in 
B. allos. On the dorsal surface of the skull the tabulars are exposed a little less 
in B. allos than they are in B. laticeps. Anteriorly the interpterygoid vacuities 
are broader in B. allos, but this difference is less between B. allos and B. laticeps 
than between B. allos and all other brachyopids. The rather long tripartite 
anterior palatal foramen in B. allos is also distinctive. 


THE SKULL OF BRACHYOPS ALLOS 
When found, the skull was lying upside down with the anterior part of the 
palate exposed. Although a large area was excavated when the skull was being 
encased in a plaster cast, no postcranial material which could positively be 
associated with the skull was recovered. In the laboratory exposed bones were 


PROCEEDINGS OF THE LINNEAN Society or NEw SoutH Wa ss, Vou. 96, Part 4 


ANNE HOWIE Diels 


glued together with Butvar B76 and impregnated with Butvar B98. Preparing 
the skull was difficult as the bone was softer in places than the matrix and the 
latter tended to remove a layer of bone as it was chipped away with an automatic 
mallet. Much of the matrix on the ventral surface of the skull roof has been 
left as a reinforcing agent as the bone is extremely thin. 


Slight dorsoventral crushing has forced the anterolateral margins of the 
skull outwards (Plate xtv) and has caused a little warping in the occipital region, 
but on the whole the skull is well preserved. 


Typical labyrinthodont ornament is present on all the skull roofing bones 

and the sensory canals where present are well impressed into this. The ornament 
is more reticular than that found in B. laticeps but is linear towards the edges of 
the bones, especially the squamosal and quadratojugal. So the suggestion by 
Bystrow (1935) that brachyopids have no regions of elongation in the skull does 
not apply to this deep cheek region where the pattern of ornament shows that the 
skull bones have grown rapidly ventrally. Watson (1956) notes this transverse 
elongation in the cheek region of Batrachosuchus, as did Save-Séderbergh (1937) 
in brachyopids in general. 
Dorsal Surface. In dorsal view the skull is very similar in shape to that of 
Brachyops laticeps except that less of the occiput is exposed. The orbits and 
choanae are rounded and the parietal foramen is small and set well back in the 
skull. A cross-section of the midline of the skull roof shows a strongly developed 
ventrally produced ridge which runs along the midline from the anterior part 
of the frontal bones posteriorly to end behind the parietal foramen (Fig. 1, x—y). 
Nine millimetres in front of this foramen the ridge divides into two, bypasses 
the foramen, and comes together again at the level of the ventral surface of the 
skull roof 6mm. behind the foramen. The result is an elongate pineal cavity 
which is loaf-shaped at the level of the external opening. 


The premaxillae, nasals, frontals, parietals, postparietals, postfrontals, post- 
orbitals, supratemporals and squamosals are very like those of Brachyops laticeps 
except that the suture between the frontals and parietals is more towards the 
front of the skull in B. allos. A flange of the nasal bone which extends lateral 
to the external naris on each side could be a septomaxilla as shown by Shishkin 
(1966) in Batrachosuchus lacer, but no nasal-septomaxillary suture is visible. 
More probably the circle of bone which floors the nostril is the septomaxilla, as 
could have been the case in Hadrokkosaurus (Welles and Estes, 1969). No 
suture can be found between the maxillary and prefrontal bones; the suture 
shown in this position in Fig. 1 is hypothetical and has been placed in its most 
likely position in relation to the ornament and the sensory canals. If it is 
correctly placed it excludes the maxilla from the orbit. A thin process of the 
jugal extends around the anterior border of the orbit to a greater extent than it 
does in other brachyopids. Much of the right quadratojugal remains and forms 
the posterolateral corner of the skull, sending sheets of bone medially to cover 
parts of the posterior and anterior faces of the quadrate. 


Ventral Surface. A ventral view of the skull shows unusually wide interpterygoid 
vacuities and an enlarged anterior palatal vacuity. The latter is tripartite 
(Fig. 2), as is the one found by Shishkin (1966) in Batrachosuchus. 


Large tusks and tusk replacement pits are found in the vomers palatines 
and right ectopterygoid, which also bears a palatal tooth. While no palatal 
teeth can be found on the vomer, an additional small tusk is present right of the 
midline just posterior to the anterior palatal vacuity. A raised median area on 
the cultriform process of the parasphenoid bears a shagreen of minute denticles 
which are not present elsewhere on the palate. 


Premaxillary, maxillary, vomer and palatine bones vary little from the 
characteristic brachyopid pattern as seen in B. laticeps. The parasphenoid body 


PROCEEDINGS OF THE LINNEAN SociEeTY oF New SoutH WaAtss, Vou. 96, Part 4 


272 A BRACHYOPID LABYRINTHODONT 


Fig. 1. Brachyops allos n.sp. a: Dorsa)] view of the skull drawn normal to the skull roof. 
b: Ventral view of the skull drawn normal to the parasphenoid. x, y: Sections of the skull 
midline at x and y. x¥%. Sensory grooves are stippled. 


PROCEEDINGS OF THE LINNEAN SocrreTy OF NEw SoutH WatgEs, VoL. 96, Part 4 


ANNE HOWIE 273 


Fig. 2. Brachyopsallosn.sp. a: Occipital view of the skull. b: Anterior view of the skull. 
ce: Lateral view of the skull drawn normal to the saggital plane of the skull. x 3. 


Fig. 3. Brachyops allos n.sp. Sketch of a section of the skull in the braincase area to show 
the (?) epipterygoid. Looking posteriorly from the proximal end of the cultiform process. x # 


PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SoutH WALES, Vou. 96, Part 4 


274. A BRACHYOPID LABYRINTHODONT 


is arched a little dorsally and extends further posteriorly than it does in B. laticeps, 
so that the exoccipitals are separated in the midline. No crista muscularis as 
noted by Welles and Estes in Hadrokkosaurus is present, but a well-developed 
ridge runs from the cultriform process behind the interpterygoid vacuity and is 
continued on the palatal ramus of the pterygoid. The cultriform process dips 
dorsally above the vomers at its anterior end but is narrower here than it is in 
B. laticeps, being clasped laterally by posterior extensions of the vomers rather 
than bearing these on its ventral surface. 


The pterygoids are typically brachyopid except in their lateral margins. 
On the left side the anterolateral border of the palatal ramus of the pterygoid 
is a true edge (edge, Fig. 2), so that on this side the ramus is less than half the 
width of the right ramus. However, a ridge on the right ramus follows a similar 
line to that taken by the true edge of the left ramus. I interpret this edge as 
being the suture line between the pterygoid and ectopterygoid, so that the 
ectopterygoid extends posterolaterally along the pterygoid. 


The right quadrate is preserved and shows an advanced level of ossification 
when compared with other brachyopids. The condyle has a lateral area and a 
larger and more ventral median area, the two being linked by a groove to form a 
screw-Shaped condyle similar to that found in other well ossified rhachitomes. 


Because of the greater degree of ossification in this specimen, an occipital 
view differs from the same view in other brachyopids in that the gap between the 
Squamosal-quadratojugal complex and the pterygoid is filled with a dorsal 
extension of the quadrate. The supraoccipital and basioccipital areas are 
unossified, but a well-developed processus lamellosus divides the supraoccipital 
space from the foramen magnum. The processus basalis of the exoccipital is 
even more fully ossified than it is in Hadrokkosaurus. A paraquadrate foramen 
is present in the right quadratojugal. The posterior face of each tabular has a 
large rugosity above the paroccipital process. 


The lamina ascendens or dorsal process of the pterygoid is much the same as 
the one present in Hadrokkosaurus except that it reaches the skull roof, touching 
the tabular medially and the squamosal more laterally. 


Epipterygoid. On the right side, 4mm. medial to and a little behind the leading 
edge of the dorsal process of the pterygoid, is a further ossification (Figs 2 (a), 3) 
which may be an epipterygoid. This originates from an 8mm. base on the 
pterygoid near its suture with the parasphenoid and ascends dorsomedially, 
becoming narrower and rounder to end just anterior to the postparietals and 7 mm. 
lateral to the midline. A small excavation beneath the anteromedial edge of the 
ascending process of the pterygoid is probably the conical recess for the basi- 
pterygoid process of the basisphenoid. Between the head of the (?) epipterygoid 
and the anterior face of the paroccipital process and attached to the latter is an 
irregular ossification which may be part of the otic capsule. 


From Husthenopteron through early amphibia like Hryops and Hdops and 
later Triassic Amphibia like Parotosaurus the epipterygoid whether more or less 
well ossified bears a contant relationship to the pterygoid ; it is found adpressed 
to the lateral (morphologically internal) surface of the dorsal process of the 
pterygoid. In Husthenopteron and early Amphibia the palatoquadrate cartilage 
‘is present as a continuous element which runs forward from the quadrate along 
the dorsolateral margin of the pterygoid, but is usually not preserved anterior 
of the epipterygoid region. In the more ‘‘ advanced ” labyrinthodonts ossifica- 
tion in the palatoquadrate is gradually reduced to a quadrate element and an 
epipterygoid element, the latter consisting of an expanded basal portion which 
usually forms the dorsal roof of the conical recess for the basipterygoid process 
of the basisphenoid, an anterodorsal process which eventually becomes the 
columella cranii of reptiles, and a posterodorsal or otic process. In Triassic 


PROCEEDINGS OF THE LINNEAN SocrETy oF NEw SoutH WALES, Vou. 96, Part 4 


ANNE HOWIE 275 


amphibians such as Lyrocephalus, Parotosaurus and Metoposaurus the otic process 
is lost. In life, no doubt a cartilaginous connection was maintained between 
the quadrate and the epipterygoid. 

Previously described brachyopids have been poorly ossified, a condition 
reflected in their fragmentary or missing quadrates and the lack of any ossification 
in the braincase area. In Brachyops allos the quadrate is better ossified than it 
is in many earlier labyrinthodonts, and it would not be unreasonable to expect 
some part of the epipterygoid to be preserved. However, the column of bone 
preserved in B. allos differs from other known epipterygoids in that its foot lies 
just medial to the pterygoid rather than being adpressed to its lateral edge, 
so that any cartilaginous connection between the epipterygoids and the quadrate 
would be extremely awkward. 

One possibility is that the element is a displaced stapes, but Watson described 
a very different stapes in Batrachosuchus, as did Bystrow (1937) in Dvinosaurus. 
Also, the bone is apparently in place within the skull and slopes dorsomedially ; 
a Stapes would be more likely to have its distal end at least outside the skull, 
and would slope dorsolaterally. 


RELATIONSHIPS OF BRACHYOPS ALLOS 


In their review of the brachyopids Welles and Estes (1969) used the method 
developed by Welles and Cosgriff (1965) for their review of the capitosaurus. 
A series of arbitrary measurements is made on the skulls, and indices are derived 
from these measurements.- The sum of the differences between indices for any 
two species is then assumed to be a useful indication of the relationship between 
these species. Welles and Estes acknowledge the various shortcomings of this 
method of analysis, one of which is the small number of specimens from which 
all measurements can be obtained. Data from additional specimens can only 
improve this situaton so the relevant figures for B. allos are tabled below. 


The figures given are for the reconstructed skull; they differ slightly from 
those obtained from the actual specimen. 


5 


Der awas 


Interorbital breadth, A : 

Breadth of skull roof across quacdratojugals, B 
Midline postorbital length, D : ie 
Length of yvomers, HE 

Midline distance between noetrile and orbits ¥ 
Breadth across vomers, G 

Height of parasphenoid, H 

Internarial breadth, J be ae ms A: at 
Length of skull roof, L .. np oe Se es ser eel 
Midline preorbital length, O : 

Distance behind orbits of parietal foramen: Pp 

Length of body of pterygoid, Q x 

Breadth across pterygoids at concavity, R 

Distance of parietal foramen in front of end skull table, T 
Midline orbital length, U . : 

Length of interpterygoid Sorat W 

Breadth of interpterygoid vacuity, Z 


Hm 


mow We bh bs Oot Pe 
Amnonwnorwowoan-.‘ ~j 


CW ob bw ow 
Or 


Indices : 
B: L . 130 U:L 19 G:E » 29 
J 20 15 D:L 51 (A) 818s 27 
A:L 41 P:A 66 Were Ib 48 
O:L 30 T:A 49 Lieve 70 


PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WaAtzEs, Vou. 96, Part 4 


276 A BRACHYOPID LABYRINTHODONT 


Sums of Differences (using Welles and Estes data as plotted in figs 2-13) : 
Brachyops allos 


aes 149 
Hadrokkosaurus bradyt 
Brachyops allos 73 
Batrachosuchus watsoni “* 
Brachyops allos 61 
Batrachosuchus brownt 
Brachyops allos 62 
Brachyops laticeps 
Brachyops laticeps 146 
Blinasaurus henwoodi 
Brachyops allos 101 
Bothriceps major 
Brachyops allos 
etd hia cece 113 
Bothriceps australis 
Brachyops allos 155 


Dvinosaurus primus 


Assuming that the smaller totals of index differences are real measures of 
relationship, this table shows that Brachyops is most closely related to Batra- 
chosuchus and next to Bothriceps—the same finding as Welles and Estes’ original 
analysis showed. The fact that Brachyops allos is closer by one point to Batra- 
chosuchus browni than to Brachyops laticeps only emphasizes the fact that until 
more brachyopid specimens are known a few points difference between two 
skulls cannot be taken as a significant measure of a relationship or a non- 
relationship ; the equivalent figures for the Brachyops allos skull before recon- 


struction indicate that B. allos is closer by five points to B. laticeps than it is to 
Batrachosuchus watsoni. 


Welles and Estes suggest that a sum of differences of 57 might reasonably 
be considered as representing a specific level. If this is so, then Brachyops allos 
should be placed in a genus separate from Brachyops laticeps and Batrachosuchus. 
This would be “ splitting ’’ in the extreme and it could be more valid to unite 
Brachyops laticens, Brachyops allos and Batrachosuchus into a single genus. 


Whether it is Brachyops or Batrachosuchus, Brachyops allos will link Welles 
and Hstes’ two Palaeogeographic units, their Australian brachyopid line, and 
their second brachyopid line which includes African and Indian-North American 
forms. 

Brachyops allos is thus a particularly well-preserved and well-ossified member 
of its family and hence shows several features not previously found in the 
Brachyopidae. These include the presence of a paraquadrate foramen, an area 
of denticles on the parasphenoid, an ossified quadrate and a probable epipterygoid. 

Being the first brachyopid to be described since Welles and Estes’ analysis 
of the family, it can be used to test their methods. These proved to be accurate 
in indicating relationships at a level a little above the generic one, but, at least 
until the discovery of more specimens allows a statistically valid index to be 


calculated, the methods cannot be used to establish generic or specific 
relationships. 


ACKNOWLEDGEMENTS 


I thank my field party, R. Evans, E. McLean and J. Warren, Mr. R. Goodwin 
who kindly allowed us to work on his property, and Professor L. C. Birch for 
having me in his department and allowing me to use that department’s station 
wagon. The work was supported initially by a Sydney University Post-Doctoral 
Research Fellowship, and secondly by a Linnean Macleay Fellowship. Subse- 
quently support has been obtained from the Australian Grants Committee. 


PROCEEDINGS OF THE LINNEAN SocrETy oF NEw SoutH WALES, Vou. 96, Part 4 


Proc. LINN. Soc. N.S.W., Vol. 96, Part 4 PLATE XIV 


Brachyops allosn.sp. a: Dorsal view of the skull. b: Ventral view of the skull. x 


clo 


Howl 


Bow arnt 


: 
; 
; 
5 


Proc. LINN. Soc. N.S.W., Vol. 96, Part 4 PLATE XV 


Brachyops allos n.sp. a: Occipital view of the skull. b: Anterior view of the skull. 
ce: Lateral view of the skull. x 2. 


a 
ni 


ve - ws 
ai | $<” ee ae eat 


_ 


is earns o 

i. quid ocak oDicte vglt ‘to wale Nang ee y 
Y, a - - = 
7” a 


ANNE HOWIE 277 


References 


BarTHotomal, A., and Howis, A. A., 1970.—Vertebrate fauna from the Lower Trias of Australia. 
Nature, Lond., 225: 1063. 

Broom, R., 1903.—On a new stegocephalian (Batrachosuchus browni) from the Karroo Beds of 
Aliwal North, South Africa. Geol. Mag., 10, 4: 499-501. 

Bystrow, A. P., 1935.—Morphologische Untersuchungen der Deckknochen des Schadels der 
Wirbeltiere. 1. Mitteilung. Schadel der Stegocephalen. Acta zool., Stockh., 16: 65-141. 

, 1938.—Dvinosaurus als neotenische Form der Stegocephalen. Acta zool., Stockh. 
19: 209-295. 

Bystrow, A. P., and Erremoy, J. A., 1940.—Benthosuchus sushkini Efr.—a labyrinthodont from 
the Eotriassic of Sharzhenga River. Trudy paleozool. Inst., 10,1: 1-152. 

CoscrirF, J. W., 1965.—A new Genus of Temnospondyli from the Triassic of Western Australia. 
J. Proc. R. Soc. West. Aust., 48: 65. 

, 1969.—Blinasaurus, a brachyopid Genus from Western Australia and New South Wales. 
J. Proc. R. Soc. West. Aust., 52: 65-88. 

Howie, A. A., 1970. A new capitosaurid labyrinthodont from East Africa. Paleontology, 
13, 2: 210-258. 

On a Queensland labyrinthodont. (In press.) 

Hux ey, T. H., 1859.—On some amphibian and reptilian remains from South Africa and Australia. 
Proc. geol. Soc., 15: 642-658. 

Matoneg, EH. J., OucErsS, F., and KrrKEGAARD, A. G., 1969.—The geology of the Duaringa and 
Saint Lawrence 1: 250,000 sheet areas, Queensland. 

Ortow, I. A., Ed., 1964.—(Elements of Paleontology) Moscow: Izdatelstov Nauka., 1-772 
(Russian). 

Owen, R., 1855.—Description of the cranium of a labyrinthodont reptile, Brachyops laticeps, 
from Mangali, Central India. Q. Jl. geol. Soc. Lond., 11: 37-39. 

Pancuen, A. L., 1959.—A new armoured amphibian from the Upper Permian of East Africa. 
Phil. Trans., 242: 207-281. 

SAVE-SODERBERGH, G., 1937.—On the dermal skulls of Lyrocephalus, Aphaneramma, and Bentho- 
suchus, labyrinthodonts from the Triassic of Spitzbergen and N. Russia. Bull. geol. Inst. 
Univ. Upsala, 27: 189-208. 

SuisHxKin, M. A., 1964.—Stereospondyli. In Orlov, 1964: 83-122. 

, 1966.—(A brachyopid labyrinthodont from the Triassic of the Russian Platform.) 
Pal. Jour., 1966, 2: 93-108. (Russian.) 

Susuxin, P. O., 1936.—Notes on the pre-Jurassic Tetrapoda from U.S.S.R. IIT. Dvinosaurus 
Amalitsky, a perennibranchiate stegocephalian from the upper Permian of North Dvina. 
Trudy Paleozool. Inst., 5: 43-91. ; 

Watson, D. M. S., 1919.—The structure, evolution and origin of the Amphibia. The “ orders ”’ 
Rhachitomi and Stereospondyli. Phil. Trans. B, 209: 1-73. 

, 1956.—The brachyopid labyrinthodonts. Bull. Br. Mus. nat. Hist. Geol., 2: 317-391. 

WELLEs, S. P., 1947.—-Vertebrates from the Upper Moenkopi formation of Northern Arizona. 
Univ. Calif. Publs Geol. Sci., 27: 241-289. 

WELLES, S. P., and Coserirr, J., 1965.—A revision of the labyrinthodont Family Capitosauridae 
and a description of Paratosaurus Peabodyi n.sp. from the Wupatki Member of the Moenkopi 
Formation of Northern Arizona. Univ. Calif. Publs Geol. Sci., 54: 1-148. 

WELLES, S. P., and Estss, R., 1969.—Hadrokkosaurus bradyi from the Upper Moenkopi Formation 
of Arizona with a review of the brachyopid labyrinthodonts. Univ. Calif. Publs Geol. Sci., 
84: 1-56. 

Woopwarzp, A. S., 1909.—On a new labyrinthcdont from the oil shale at Airly. Rec. Geol. Surv. 
N.S.W., 8: 317-319. 


Abbreviations 
er .. conical recess for the basipterygoid Pp .. parietal 
process of the basisphenoid pa .. palatine 
dpt .. dorsal process of the pterygoid pf .. postfrontal 
ect .. ectopterygoid pm .. premaxilla 
edge .._ true edge on the palatal ramus of po _.. _ postorbital 
the pterygoid pp .. postparietal 
eo .. exoccipital prf .. prefrontal 
ept .. epipterygoid psp .. parasphenoid 
f .. frontal pt .. pterygoid 
j jugal q .. quadrate 
m maxilla qj -.- quadratojugal 
n nasal sq .. squamosal 
ot .. otic ossification t .. tabular 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WaAtLgEs, Vou. 96, Part 4 


THE DEPOSITIONAL ENVIRONMENT OF THE ORGANIC DEPOSITS 
ON THE FORESHORE AT NORTH DEEWHY, NEW SOUTH WALES 


A. R. H. MARTIN 
School of Biological Sciences, University of Sydney, Sydney, N.S.W. 


[Accepted for publication 21st July 1971] 


Synopsis 
Pollen analysis of the coastal site at North Deewhy necessitates qualification of a previously 
published statement that could be taken to imply that the peat at this site is of brackish or marine 
origin, and that it is ipso facto evidence of a higher-than-present sea level stand less than 4,000 
years ago. 


The so-called ‘‘ peat ’’, exposed on the foreshore at North Deewhy (southern 
side of Long Reef, lat. 33° 44’S., long. 151° 19-7’ K.), has attracted attention 
for many years and was the subject of a radiocarbon dating (Hubbs, Bien and 
Suess, 1963). 


I was asked in 1962 to pollen-analyse a sample from this deposit to find 
whether marine influence was indicated. The position of this sample in the 
Sequence was apparently at the same level as the C,,-dated sample, i.e. the base 
of the organic deposits, just above heavy blue clay. The analysis showed high 
Casuarina values (Casuarina is only dominant at the base of the sequence), 
rather high Chenopodiaceae values and the presence of Hystrichosphaeridae, 
which were then generally regarded as brackish and marine indicator organisms. 
On this basis, it was concluded that the sample did show some evidence of 
proximity to the sea. 


The C,, sample was subsequently published (loc. cit., p. 259, sample LJ451), 
the date being 3980+150 yrs. (2030 B.c.), and bore the comment ‘‘ The peat 
is possibly indicative of a higher-than-present sea-stand ; pollen analysis indicates 
that it is probably of brackish or saltwater origin’’. This comment goes further 
than the data available warranted at the time, since peat of saltwater origin in 
this latitude would almost necessarily be a mangrove peat, and none of the 
palynological evidence indicated mangrove presence. Except for the Hystri- 
chosphaeridae, the aquatic plants and algae identified were freshwater forms, 
though some were euryhaline. The determination of the depositional environ- 
ment of this peat is crucial to its use as evidence of sea-level change, since, if the 
peat is of salt or brackish water origin, its position must indicate a sea-level 
higher than to-day’s, whereas if it is essentially a freshwater environment (even 
if influenced by proximity to the sea) it can offer evidence only of a negative kind. 


A pollen analysis of the entire short vertical sequence is now available. 
Fig. 1 (a) represents the pollen of woody plants and herbaceous plants of 
apparently non-local origin. Fig. 1 (b) represents aquatic and swamp plants 
and other locally produced herbs. By ‘local’ is meant those plants most 
likely to have been floristic components of the actual area of deposition at the 
time. These latter are expressed as a percentage of the pollen sum formed by 
the elements of Fig. 1 (a). The stratigraphic sequence at the point of sampling 
is depicted on the right-hand side of Fig. 1 (a) and is as follows : 


0 cm. upwards .. Dune sand with plant roots 
(arbitrary datum) 
Uf Gil Gm, .. Sandy limnic humus, more peaty at base 
43cem.-l7cem. .. .. Humified and compressed coarse detritus mud 
63 cm.—43 cm... .. Compressed grey-brown fine detritus mud 
Below 63cm. .. .. Stiff blue-grey clay. 


PROCEEDINGS OF THE LINNEAN Society or NEw SoutH WALES, Vou. 96, Part 4 


A. RB. H. MARTIN 279 


At an undetermined depth the blue clay appears to be resting on eroded brick 
earth (laterite), which outcrops seawards and can also be seen dipping under the 
deposits farther east along the shore. 

The pollen diagrams, from bottom to top, show: 

(i) Shallow lacustrine conditions with Chara and Pediastrum, Triglochin 
(or Potamogeton), Cyperaceae and probably Casuarina fringing woodland. 

(ii) Casuarina being replaced by Eucalyptoideae: declining Chara, but 
desmids (Pleurotaenium and Cosmarium) abundant; Dinoflagellates and Hystri- 
chosphaeres becoming commoner. 


NORTH DEEWHY (LONG REEF). & 
ee $ 


a gf Kae? o SS 08 
ios oe ners » 5 OF SOKO LS 
LO & WRG Re OL PERS EF 2 
we of Ronco wy? WS ONG EAM & OG 
V y 4 
oe Qe SR oR MOSS ST SF PE 
C66 Cf (COne af 
! 


») 
Y Cc 
°F, 4 . 
ent 20°%o / , 20 ee i 60 Ce te 20 oO 60 YOe G lO ¢ 
{e) 


CARPO- IGULIFLORAE 
Sf BROTUS wr V 


} 


' 
SI 
+) 
ELAEOCARPUS cf BAUERA HIBBERTIA 
+) ab ob ic ) b 
OLEACEAE mason 
©. LACKAMA BREYNA b | » ) 
» % > 980 +150 
‘ ry ~~ = 
ir i= Prana rr ire Pie eerie eee ie rie esese 


Soe pn Cpa gg 
() 20°%o 


Fig. la. Pollen diagram of deposits at North Deewhy: Arboreal and herbaceous pollen 


other than local and swamp plants. 
The Angophora — bloodwood eucalypt type is shown (white) as a component of the total 


Eucalyptus-Angophora pollen percentage. 


NORTH DEEWHY (LONG REEF) d as 
. & 
re Ss 
o x 
. 2 
© 
CCC ay ZO Ge 
~ =~ - - ---TRIGIES%e 
= SCIRPUS TYPE 
ez 72 F=TSi90% 273°%e 
INOFLAGELLATES 
+ 
& rb. ierana ee 
EV ge gs Boose UD at es ers Vpn er oe Uc oad et et Noe en re Ae e O es Ve Th i Vd Kee] ld ivi Up Us Ms 0 sD con corn Be Ber ed ea Vc Lt ae De Tied Cee A Bpeet Ce el SSS TE 


Gales 
fo) 20% 
Fig. 1b. Pollen diagram of deposits at North Deewhy : Local (i.e. presumed components of 


the peat) flora, expressed as a percentage of pollen sum of Fig. la. Cyathea and trilete spores 
other than Gleichenia are percentaged with the non-local flora, but are included with other 


Pteridcphytes for convenience. 


(iii) The fine detritus mud changing quite sharply to a coarse detritus mud ; 
much higher values for Leptospermoideae (Leptospermum, Melaleuca and Kunzea 
not being easily separable) probably indicating nearby ti-tree swamp, but wood 
fragments occurring neither at the point of sampling or elsewhere along the 
exposure. The influx of Leptospermoideae is probably responsible for an 
artificial lowering of the background pollen of Eucalyptus, ete. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Watss, Von. 96, Part 4 


280 DEPOSITIONAL ENVIRONMENT OF ORGANIC DEPOSITS, DEEWHY 


(iv) Peat development, with occasional very compressed Cyperaceae 
rhizomes, at about the point at which Cyperaceae pollen becomes abundant in 
the profile. This level also corresponds to a step cut in the deposit which is 
apparently due to a difference in erodability of the two sediment types. 

(v) The peat grading into the more limnic sandy-humic layer upwards, 
itself not sharply demarcated from the overlying sand, but apparently water- 
logged conditions prevailed to the time of obliteration of the swamp, as indicated 
by the vast quantities of Triglochin and Cyperaceae pollen and the entry of 
Philydrum and Nymphoides into the profile. Leptospermoideae, apart from 
Callistemon, which actually increases slightly, decline towards the top of the 
profile, resulting in a recovery of the non-local Casuarina, Eucalyptoideae and 
Callitris. 

It is now clear that the lower third of the sequence represents a lake or 
freshwater lagoon with a desmid and Pediastrum phytoplankton. There is no 
warrant for the belief that the deposit was in a marine or brackish (as that word 
is usually understood) environment, and the presence of ferrous sulphide (FeS,) 
erystals, which occur in the basal lake mud, is quite compatible with formation 
in a freshwater lagoon under slight marine influence (e.g. Degens, 1965). The 
freshwater tolerance of hystrichosphaeres (in this context, microplanktonic 
fossils with spherical bodies and radiating processes that are not obvious Dino- 
flagellates) appears to be higher than formerly supposed (Churchill and Sarjeant, 
1962), and in the general context of this deposit they can hardly be said to provide 
clear evidence of high salinity. 

The central part of the sequence, which lacks even equivocal indicators of 
salinity, must have been produced under waterlogged or near-waterlogged 
conditions if not open water, because of the dominance of swamp-living angio- 
sperms. A slight rise in water level may be implied by the replacement of a 
Leptospermoideae swamp by a Cyperaceae-Nymphoides-Triglochin swamp in 
the upper part of the sequence, but the absence of any wood fragments makes 
the precise meaning of the high Leptospermoideae values uncertain. The 
changes in flora indicated by the pollen analysis do not reflect a straightforward 
plant succession and could have been brought about by minor changes in water 
level, which itself would be influenced by factors such as the movements of nearby 
dunes, shoreline recession and migration of the mouth of the nearby Deewhy 
Lagoon. 

An opportunity to obtain the true level of the deposit did not occur until 
this year. Consequently, the precise position from which the samples were taken 
could no longer be located, though the general position was located within a 
few metres. The outcrop along the beach is stratigraphically uniform and fairly 
flat except for a slight rise at the eastern edge. Thus, the level obtained is 
believed to be representative of the outcrop as a whole. At the new point of 
measurement the surface of the deposit is 3-0 m. above standard datum (N.S.W. 
standard datum is 2-93 ft. or 0-89 m. above Indian Springs Low Water at Fort 
Denison, Sydney). The tidal range at Narrabeen is approximately the same as 
at Fort Denison, i.e. 0 to +2-0m., and the clay/mud transition is 2-2 m. above 
standard datum, the deposit being 80 cm. thick. A second measurement a few 
metres from the first gave a thickness of 75 cm., compared with 64 cm. at the 
point of sampling in 1965 when the pollen samples were collected. 

Allowing for the compression which has taken place during the burial of 
the sediments under dune sand, the deposit appears to have formed within 
about 2:3m. of local HHWS. The water level in the existing nearby Deewhy 
Lagoon was 0-9 m. below the peat base at the time of survey and peat forming 
at the present day round the margin of the lagoon is approximately 0-6 m. 
below the base of the peat on the foreshore. One can point to a number of still 
existing freshwater peat swamps, at levels ranging from several metres higher 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vou. 96, Part 4 


A. R. H. MARTIN 281 
to about 0-5 m. lower than the North Deewhy deposits, in similar areas along the 
central coast of New South Wales (e.g. Kurnell Peninsula). In each case local 
water-table conditions determine the level at which peat formation occurs. 
Needless to say, such deposits cast very limited light on existing sea level. 


An interesting sidelight of the work is the finding of a few pollen grains of 
several unexpected plants, e.g. Nothofagus, Rhodomyrtus and cf. Ackama. The 
two latter seem to represent a coastal rainforest or wet sclerophyll element in 
the flora. Rainforest remnants still grow on the Narrabeen shales of the Palm 
Beach Peninsula, e.g. at Bilgola, a few miles to the north of Deewhy, though 
neither of these genera are now present locally. Nothofagus, on the other hand, 
appears to represent a long-range component. The grain is of the N. moorei 
type, the nearest source of which, today, would be the Barrington Tops forests, 
ca. 200 km. distant. The single grain is unlikely to be a contaminant, as it 
contains small ferrous sulphide crystals, as do other pollen grains at this level. 


ACKNOWLEDGEMENTS 


I am indebted to the Sydney University Research Committee and to the 
Nuffield Foundation for grants for palynological research. Thanks are due to 
Dr. M. Williams and to N. P. Thiem for survey data, and to the N.S.W. 
Department of Lands for information and maps relating to the height and 
location of benchmarks. 


References 

CHURCHILL, D. M., and SarRJEANT, W. A. S8., 1962.—Fossil Dinoflagellates and Hystrichosphaeres 
in Australian freshwater deposits. Nat. (Lond.), 194: 1094. 

Decens, E. T., 1965.—Geochemistry of Sediments. Prentice-Hall, N.J. 

Housss, C. L., Bren, G. S., and Susss, H. E., 1963.—La Jolla Natural Radiocarbon Measurements. 
Ill. Radiocarbon, 5: 254-272. 

Tuom, B. G., Hains, J. R., and Martin, A. R. H., 1969.—Radiocarbon evidence against higher 
postglacial sea levels in Eastern Australia. Marine Geol., 7: 161-168. 


PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WALES, Vor. 96, Part 4 


PETROLOGY AND STRATIGRAPHY OF THE BRAYTON DISTRICT, 
NEW SOUTH WALES 


SUZANNE Y. O’ REILLY 
Department of Geology and Geophysics, University of Sydney 


(Communicated by Dr. R. E. Wass) 
[Accepted for publication 23rd June 1971] 


(Plate Xv1) 


Synopsis 

An angular unconformity separating Upper Ordovician beds from (?) Silurian sediments 
and basic and acid volcanics has been mapped in the Brayton area. It is shown that the fine- 
grained silicic igneous rocks constitute a series of contemporaneous volcanic flows within the 
(?) Silurian succession, a conclusion at variance with earlier opinion. These volcanic rocks and 
the Marulan batholithic rocks are therefore distinct and separate entities. The hybrid zones of 
previous workers are re-interpreted as zones of increasing metamorphic grade where intrusion of 
granitic rocks into the silicic voleanics produced a contact aureole. A Tertiary dolerite body and 


two Tertiary basaltic flows exhibit closely correlative mineralogy, while the dolerite shows limited 
upward differentiation. 


INTRODUCTION 


The Brayton area lies approximately 130 miles south-west of Sydney, 10 
miles north of Marulan. About 12 square miles were mapped in detail as shown 
may 10m, IL 

Upper Ordovician slates, quartzites and phyllites have undergone much 
regional deformation, culminating in an isoclinally folded sequence with a 
macroscopic meridional trend and approximately vertical dips. The (?) Silurian 
succession, which includes three distinct units, overlies the Ordovician rocks with 
angular unconformity. At the base, a series of shales, sandstones and calcareous 
siltstones with a general north-south strike and steep westward dip outcrop. 
Succeeding these sediments are basic volcanic flows with rare sedimentary lenses 
and a small limestone pod. Extensive silicic volcanic flows ranging from 
toseanites to dacites with some tuff bands comprise the youngest unit. 


Field and petrological evidence indicates that the (?) Silurian sediments, 
basic voleanies and silicic volcanics constitute a conformable sequence and have 
shown synchronous response to low-grade metamorphism and _ regional 
deformation superimposed on the already deformed Ordovician sequence. 


Granitic rocks forming part of the Marulan batholithic complex have intruded 
these Palaeozoic strata, producing contact metamorphic effects within an aureole 
at least one mile wide. An intermediate dyke of unknown age intrudes one of 
the granitic bodies. 


The remaining rocks are Tertiary in age. These include the Billyrambija 


dolerite, two basalt flows, and the associated Tertiary sediments, which are 
rich in a fossil flora. 


PREVIOUS WORKERS 
First geological observations made in the area were those of Clarke (1845). 
He stated that the Wollondilly River from its source to the junction with Paddy’s 
River cut through igneous and metamorphic rocks which were laid bare over a 
considerable area between the Cookbundoon and the Uringalla. 


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SUZANNE Y. O'REILLY 283 


Craft (1928) recorded that the Lower Palaeozoic rocks of the Wollondilly 
area are mostly Silurian and Devonian and ‘‘... consist of quartzites, slates 
and claystones together with some shales and intrusive granites. The general 
strike is about 10° east of north...”’. 

Woolnough (1909) investigated the general geology of the Marulan-Tallong 
district, to the south of Brayton. More detailed petrological studies on the 
Marulan batholithic rocks at South Marulan were carried out by Osborne (1948). 
Lovering (1950), Svenson (1950) and Osborne and Lovering (1952) studied in 
detail the ‘“‘ quartz porphyrites’”’ and ‘‘ quartz porphyrite hybrids’ at South 
Marulan. These rocks are similar to the silicic voleanics at Brayton. 


LEGEND 
Tertiary sandstone. | GEOLOGICAL MAP 


Tertiary Billyrambija dolerite. 
SSS Tertiary basalt. 
Lockyersleigh adamellite. 
Towrang granodiorite. 
Brayton volcanics, 
Silurian zeae Limestone pod. 
cal 


OF THE HA 


/ 
BRAYTON WATT 
N.S: 


pevonien| 


Longreach volcanics. 


EZ Wollondilly beds. 


eee SS 
onde cien [LIL] Ordovician metamorphics . “ih oND ILL S, 
Basic Dyke. w seae 
Sear Aplite Dykes and Pods. 
Geological Boundary -Accurate. ‘ / 
Da 
x 


Geological Boundary -Inferred. 


Strike and Dip Reading-Overturned, 


= River. 


Creeks. 


Roads. 


Trig. Station 


Homesteads. 


Railway Line. 


In a series of papers, Naylor (1935a, 1935), 1938, 1939 and 1949) outlned 
the general geology of the Goulburn district. Of these, the first and last papers 
are particularly pertinent as rocks of the Brayton area are mentioned. Ordovician 
and Silurian boundaries, the extent of the Marulan Batholith and Tertiary 
basic igneous rocks were mapped on a scale of four miles to an inch. No differ- 
entiation into formations was effected, while the silicic voleanics were included 
as an integral part of the Marulan Batholith. 


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284 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


Browne (1933) first mentioned the dolerite outcropping at Billyrambija 
Trig., near Brayton. This he considered to represent a denuded Tertiary sill. 


ORDOVICIAN ROCKS 
The Ordovician strata are typically highly deformed, have a regional 
meridional trend, are steeply dipping westward, and contain an Upper Ordovician 
eraptolite fauna. 


Primary bedding is defined by alternating layers of dark blue-grey 
carbonaceous slates and quartzites, both ranging typically from five to 12 feet 
in thickness. Within the quartzite layers are occasional] fine bands of phyllitic 
siltstones six inches to 12 inches in width. 


Structure: These beds probably lie on the overturned limb of a macroscopic 
fold or fold system which also involves the (?) Silurian sequence. Beds are 
almost vertical and all foliations measured are steeply dipping. Two fold types 
are evident on a mesoscopic scale. The first comprises similar folds with 
attenuated limbs and wider hinge areas. Wavelength is about 10 feet, with a 
comparable amplitude, the strike of the axial plane being approximately east-west. 
The axis is vertical ; the fold surface is primary bedding. The second comprises 
small isoclinal folds with average wavelength approximately one foot and 
amplitude of two to four inches. The axial plane is vertical and strikes east-west, 
while axes plunge steeply westward. 


Slates exhibit greater deformation than the more competent quartzites. 
The degree and style of deformation are not constant throughout the sequence. 


Two further penetrative structural elements occur. One is a mineral 
streaking, approximately axial plane for the small isoclinal folds. The other 
is a layering defined by lenticular pods and isolated small closures. These show 
somewhat random orientation, but an overall east-west trend is evident. 


Age and fauna: The dark grey carbonaceous slates yield two graptolite forms, 
Climacograptus bicornis and Dicellograptus sp. An age of Upper Gisbornian to 
Bolindian may thus be deduced (Thomas, 1960). Naylor (1949) previously 
recognized Climacograptus bicornis, Diplograptus quadrimucronatus and Dicello- 
graptus complanatus in the vicinity of Brayton, but no precise locality was given. 
Petrography: The quartzites consist of detrital quartz grains with rare felspar 
laths in a matrix of sericite and chlorite. Characteristic heavy minerals are 
smoky blue to blue-green pleochroic tourmaline, brownish-yellow to olive-brown 
pleochroic hornblende and opaque grains. 


The matrix, forming up to 40% of the rock, has an incipient microscopic 
layering. Quartz shows evidence of deformation, including undulose extinction, 
Bohm lamellae and growth of small strain-free grains at sub-grain boundaries. 
Strings of tiny inclusions as well as apatite and zircon needles indicate a possible 
igneous origin for at least some of the quartz. 


The phyllitic siltstones are compositionally equivalent to the quartzites. 
Macroscopically, the sheen and crenulations on penetrative surfaces are those of 
true phyllites. 


The slates contain quartz (50%), sericite (20°%), carbonaceous matter (18%) 
and opaque granules with rare zircon and tourmaline. 


Contact metamorphism: The metamorphic grade attained by the pelitic rocks 
close to the granitic contact appears to be consistent with that of the hornblende 
hornfels facies. Quartz-muscovite-biotite-cordierite is the characteristic 
assemblage. The carbonaceous slates show development of andalusite needles 
up to 4mm. long. Retrogression of the andalusite to sericite aggregates is 
ubiquitous. 


PROCEEDINGS OF THE LINNEAN SociETy oF NEw SoutH WALES, VOL. 96, Part 4 


SUZANNE Y. O’REILLY 285 


(2) SILURIAN SEQUENCE 

As the age of this sequence cannot be strictly defined, it is referred to as 
(?) Silurian thus. The (?) Silurian beds are separated from the Ordovician 
rocks by a distinct angular unconformity which is discussed in detail below. 
Three units may be distinguished by lithological differences. These are the 
Wollondilly beds, Longreach volcanics and Brayton volcanics in ascending 
stratigraphic order. Their conformity is indicated by structural features 
including close correspondence of dips, strikes and orientation of the two 
ubiquitous joint sets. All (?) Silurian rocks show comparable mineralogical 
adjustments to a low-grade metamorphic environment. Thus synchronous 
deformation may be inferred. 


WOLLONDILLY BEDS 
A measured section extending from the unconformity against Ordovician 
rocks (29387150)* to the top of the Longreach volcanics (29497150) indicates 
the thickness of this unit is approximately 2,800 feet. 


Four hundred feet of calcareous siltstones containing a brachiopod fauna 
succeed a thin conglomerate band (about five feet thick) which defines the base 
of the sequence. ‘The remainder consists of interbedded sandstones and shales. 
Two prominent joint sets of consistent orientation dominate these rocks and 
bedding is often discernible only as a gross lithological layering. A penetrative 
cleavage is developed in the fine shale bands. 


The Wollondilly beds and Ordovician rocks are unconformable. Because 
of overturning, the Ordovician overlies the (?) Silurian in the field. Angular 
unconformity is based on the following criteria : 


(1) The two distinctive faunas indicate an hiatus at least from Upper 
Ordovician to possibly Upper Llandovery. 

(2) The lithologies and structural complexity of the Ordovician rocks which 
must have suffered regional deformation before (?) Silurian sedimentation 
are distinctive. 

(3) The trend of the boundary is in places transgressive to the strike of the 
(2?) Silurian strata and suggests a gently undulating depositional surface. 

(4) A conglomerate bed at the base of the (?) Silurian sequence contains 
pebbles from the Ordovician succession. 


Sedimentary structures including current bedding, small-scale minor graded 
bedding, load casts and trace fossils indicate the sequence has been overturned. 
That bedding dips more steeply than the dominant cleavage, if this is axial plane 
cleavage, also supports overturning. 


Age and fauna: The calcareous siltstones contain a profuse brachiopod fauna : 
fossils are abundant parallel to original bedding planes. Preservation is extremely 
poor but three forms were identified. 


The first is characterized by ornamentation typical of Atrypa. Absence of 
further distinctive features precludes determination. The second is a delthyrid 
brachiopod, internal moulds of pedicle valves being preserved. Thus a maximum 
time span from Lower Silurian to Middle Devonian is indicated. This brachiopod 
could be Hovwellella sp., which would refine the time interval to from Upper 
Llandovery to Gedinnian. However, identification is not unequivocal due to 
the absence of definitive brachial valves. The remaining brachiopod occurs as 
internal moulds of brachial valves and has features consistent with those of 
Schuchertella sp., which is a Lower Devonian form (N. M. Savage, pers. comm.). 

The unconformity between Ordovician and (?) Silurian beds is thus firmly 
established on faunal grounds. Because generic attribution cannot be positively 
made, the sediments containing the brachiopod fauna are tentatively called 
(7) Silurian. 

* Grid references refer to the Goulburn I : 250,000 sheet SI 55-12. 
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2386 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


Naylor (1949) considered these beds to be Lower Silurian, based on the 
lithological similarity with the Jerrara Beds near Bungonia, which yields a Lower 
Silurian graptolite fauna. He assigned an Upper Silurian age to the limestone 
pod in the Longreach volcanics because of the profusion of Upper Silurian 
limestones in the Goulburn district. 


Petrography: The calcareous siltstones consist of approximately 30° carbonate, 
30°% quartz, 20% plagioclase and 20%, interstitial chloride. Subangular quartz 
eranules are generally less than 0-:1mm. across, and show serrated edges. 
Plagioclase laths up to 0-3 mm. long are An)_; where unaltered to chlorite and 
zeolitic minerals. Carbonate occurs as granular aggregates possibly detrital 
in origin and as patches up to 0-5mm. across, apparently replacing original 
matrix. Authigenic chlorite occurs as a cement. Where developed, massive 
or spherulitic aggregates of chlorite randomly replace plagioclase and original 
matrix. The chlorite is pale green, non-pleochroic and isotropic. 

Detritus and matrix of the sandstones and shales are identical, only grain- 
size being distinct. Quartz, felspar, zircon and hornblende, tourmaline and 
opaque grains comprise the detritus. Grains are subangular and sorting is fair. 

Quartz (60° to 70°) is of two types: angular grains with sharp extinction 
and subangular grains with undulose extinction and sometimes showing sub- 
grain development. These grains also contain strings of tiny particles and 
acicular zircon crystals. Marginal serration of all quartz is typical. 


Felspar ranges from 5% to 20% in mode and is extensively altered to sericite 
and chlorite. Some show vestiges of multiple twinning but composition cannot 
be determined due to alteration. Interstitial material includes abundant sericite 
as fibrous aggregates which often encroach on detrital plagioclase margins. A 
colourless or pale green isotropic chlorite is subordinate. This cement constitutes 
up to 20% of the rock. 


Contact metamorphism: The metamorphic rocks may be divided broadly into 
lower and higher grade assemblages dependent on distance from the granitic 
contact. Only original sandstones and shales outcrop within the aureole and 
observed assemblages are as follows : 
(1) Lower grade : 
Quartz-albite-muscovite-biotite 
Quartz-muscovite-chlorite-opaque minerals 
(2) Higher grade : 
Quartz-muscovite-cordierite-biotite 
K-felspar-biotite-plagioclase-corundum. 


Different assemblages probably reflect the original composition of the 
sediment, although no silica-poor rocks were found outside the aureole. Lower 
grade rocks are consistent with parageneses of the albite-epidote hornfels facies, 
those of higher grade rocks generally le in the hornblende hornfels facies. 
However, where potassium felspar is associated with corundum, pyroxene 
hornfels facies may be indicated. 


LONGREACH VOLCANICS 


The Longreach volcanics comprise a series of basaltic flows best exposed at 
29567088. Outcrops extend over about half a mile between the Wollondilly 
beds and the Brayton volcanics. As flows are now vertical, this represents the 
thickness of the sequence. Small bands of sediment occur rarely between these 
flows and a small limestone pod outcrops at 29637092. Trend of flows is 
meridional, while two sets of prominent vertical joints are developed with 
orientations consistently 290° and 245°, as in the Wollondilly beds. 


At 29557057 basaltic dykes transect the Wollondilly beds. These intrusions 
may represent feeder dykes for the flows of the Longreach volcanics as they show 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WaAteEs, Vou. 96, Part 4 


SUZANNE Y. O’REILLY 287 


analogous low-grade alteration. Dykes show no preferred orientation and thus 
were not controlled by any pre-existing structural elements. 


Petrography: The basalts are extensively altered and similar to spilitic basalts 
in fabric, mineralogy, chemistry and geological occurrence—the four criteria 
discussed by Vallance (1960). Three distinct types of alteration are evident, 
based on the dominant phases: (a) chlorite-epidote-tremolite, (6) chlorite, and 
(c) prehnite-calcite. Mutual gradations exist with no systematic distribution 
of alteration types. 

The basalts generally show an intersertal texture with glomeroporphyritic 
aggregates of augite, plagioclase or augite and plagioclase. Flow foliation is 
evident but often obscured by alteration ; amygdales abound. A subvariolitic 
texture is common (e.g. 33692*). Quartz xenocrysts are abundant; these 
generally show no mantling by pyroxene or evidence of reaction. 


Plagioclase occurs as microlites, laths and glomeroporphyritic aggregates 
of laths up to 1 mm. long, with composition about An;. Swallow-tail terminations 
indicate rapid chilling. Relict augite crystals have 2Vx=60°, Z\c=40° and are 
colourless. Subhedral grains range from 0:5 mm. to 2:0 mm. across; glomero- 
porphyritic aggregates are common. The original mesostasis has been almost 
completely obliterated. It was probably glassy to cryptocrystalline ; 33692 
Shows glass shards pseudomorphed by chlorite. Felspar microlites, tremolite 
needles, turbid sphene granules, chlorite and a felted layer silicate now constitute 
the groundmass. 


A typical chlorite-tremolite-epidote assemblage is present in 33694. Hpidote 
occurs aS granular aggregates, prismatic crystals or sheaves of bladed crystals. 
Colourless granules replace plagioclase, but most epidote lines or fills amygdales. 
Optical properties are highly variable, often within the one crystal, some being 
pleochroic from bright to pale yellow with 2Vx—70°, some being colourless with 
2V,—15°. Two types of chlorite are present. One is colourless and isotropic, 
the other pleochroic from pale blue-green to pale yellow-green and shows brilliant 
anomalous blue interference colours. The chlorite generally replaces augite 
but may also infill or line amygdales. Tremolite occurs as felted aggregates of 
acicular needles (ca. 0-2 mm. long) in amygdales, as tiny needles (less than 
0-1 mm. long) in the groundmass and as large rods associated with altered augite. 
It is generally colourless, but larger laths may be faintly pleochroic from colourless 
to pale blue-green. 2Vx is about 80°, ZAc is about 20°. Rare prehnite occurs 
as spherulites replacing augite and groundmass or infilling amygdales. 


Dominant chloritic alteration is shown in 33687 and 33682. Relict augite 
and original glass shards (e.g. 33692-3) are altered to pale green chlorite showing 
brilliant purple anomalous interference colours. Amygdales are typically infilled 
with calcite or quartz, with or without chlorite. Felspar laths are extensively 
sericitized. 


Prehnite-calcite alteration is typified by 33690-1. Here felspar shows no 
clouding. Augite is much altered to radiating sheaves of prehnite closely 
associated with patches of chlorite. Prehnite is optically positive with 2V ranging 
from 40° to 60° within a crystal. A pinkish hue in plane polarized light may be 
due to included haematite. Prehnite occurs as spherulites about 0-3 mm. in 
diameter, replacing augite and groundmass non-selectively. It also occurs as 
Massive or sheaf prehnite where it pseudomorphs phenocrysts, appears not 
to replace the groundmass and is invariably rimmed by opaque grains. Abundant 
calcite is intimately associated with sheaves of prehnite and also occurs as ragged 
anhedra up to 3mm. across and as amygdale infillings. Tremolite, sphene and 
epidote are present in the groundmass. Small orange spherulites are dotted 


* Five-figure numbers used throughout the text refer to rocks in the Sydney University 
Geology Department collection. 


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288 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


sparsely throughout the prehnite. Amygdale assemblages include laumontite (?)- 
prehnite-calcite and quartz-calcite-chlorite-opaque minerals. 


Contact metamorphics: In the lower grade rocks (e.g. 33701, 33709, 33711) the 
characteristic assemblage includes tremolite-actinolite, plagioclase, calcite, 
quartz, minor sphene and opaque grains. Plagioclase-actinolite-calcite 
assemblages appear to be analogous to patches of prehnite-epidote-chlorite in 
rocks outside the aureole. Plagioclase (ca. An,,) occurs as granoblastic 
aggregates of anhedral grains and as tiny laths in the groundmass. Tremolite- 
actinolite pseudomorphs original glomeroporphyritic pyroxene and occurs as a 
felt of tiny laths and needles replacing the original mesostasis. It is pleochroic 
with X=pale yellow-green, Y=colourless, Z=pale blue-green. 2Vx is 80°: 
ZAc ranges from 15° to 20°. Refractive index @ is 1-620. 

The higher grade rocks develop two distinctive types of assemblages, one 
abundant in pyroxene, the other dominated by hornblende and plagioclase. 
Both types still retain features of the original basaltic fabric. The texture is 
invariably blastoporphyritic ; original amygdales and quartz xenocrysts are 
recognizable, although both are recrystallized and the former have mineralogically 
adjusted. The patchy, variolitic aspect of the flows is also preserved. 


Hornblende-plagioclase rocks are the more abundant. The mineral 
assemblage comprises hornblende, plagioclase, quartz, sphene, epidote, opaques, 
potassium felspar and sphene. Hornblende pseudomorphs original augite 
glomeroporphs, occurs as discrete rods up to 2 mm. long and forms a groundmass 
network. Pleochroism with X=yellow, Y=bright yellow-green, Z—blue-green 
is distinctive. 2Vx is high, ca. 85°, Zc is ca. 20° to 24°. Retrogression to a 
yellow uralitic product ranges from incipient to extensive. Plagioclase, An,, 
to An,, occurs as euhedral laths up to 0-5 mm. long. Pyroxene-rich assemblages 
occur as veins or patches in the normal hornblende-plagioclase rocks. Minerals 
present are diopside, rare hedenbergite, plagioclase (An;,), calcite, scapolite, 
hornblende, sphene, garnet, epidote and rare opaque granules. 


BRAYTON VOLCANICS 


The youngest unit within the (?) Silurian succession represents a period of 
silicic voleanic activity. Toscanites, dacites and tuffs, which extend beyond 
the area mapped, outcrop over about one mile normal to strike. As the sequence 
is now vertical, this represents a thickness of 5,000 feet. Evidence for vertical 
tilting is based on (a) marked variation in rocks types on a foot scale strati- 
graphically, while a few distinctive flow types may be traced for over a mile in 
a north-south direction and (b) the apparent conformity of all the (?) Silurian 
rocks. 


Age and geological relationships: The silicic voleanics were extruded prior to 
emplacement of the Marulan batholithic rocks. Contact of the Brayton volcanics 
with the granitic rocks is sharply delineated and intrusion by the Lockyersleigh 
adamellite is undoubted, as evidenced by the metamorphic effects discussed 
below. Four factors are most significant : 


(1) These rocks are extrusive as is first indicated by their conformity with 
the Wollondilly beds and the Longreach volcanics. The Brayton 
volcanics form an integral part of the stratigraphy. 


(2) Recognition of flow units, presence of tuff bands, flow lineation (Plate 
XVI, fig. 1) and relict glass shards (Plate xvi, fig. 2) indicate a volcanic 
origin. Evidence for extrusion is observed in several exposures whose 
brecciated nature indicates explosive activity. Angular toscanite 
fragments (1 in. to 6 in. across) are separated by a network of essentially 
epidote-hornblende domains up to1 in. in width. This network probably 
represents original glass which formed a groundmass for the breccia. 


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SUZANNE Y. O'REILLY 2389 


Subsequent low grade alteration effected chemical redistribution involving 
both glass and breccia fragments. 
The intrusive nature of the granitic rocks demonstrates the older age of 
the Brayton volcanics. That considerable time intervenes is a logical 
result of the conformity of the whole (?) Silurian sequence. A period of 
structural deformation and low grade rock alteration must have occurred 
before introduction of the granitic rocks. As discussed above, 
the Brayton volcanics are tentatively assigned to the Upper 
Silurian although a Lower Devonian age is not precluded. The age of 
the Marulan Batholith on present data may be pre-Upper Devonian 
to Carboniferous (see below). 
(4) Chemical evidence is inconclusive as the two chemical analyses available 
(Osborne and Lovering, 1952) demand no genetic relationship between 
the silicic volcanics and granitic rocks. 


In summary, the extrusive nature and approximate time of extrusion indicate 
that the Brayton volcanics and the Marulan batholithic rocks are discrete entities. 


This departs from the concepts of previous workers, notably Woolnough 
(1909), Naylor (1935a) and Osborne and Lovering (1952), who considered the 
silicic volcanics to be shallow intrusions intimately associated in time and possibly 
comagmatic with rocks of the Marulan Batholith. 


Petrography: Mesoscopically the silicic volcanics are fine to medium grained 
and range from dark grey green to greenish blue; flow foliation is sometimes 
visible. Phenocrysts of euhedral quartz, zoned plagioclase, alkali felspar, 
hornblende and biotite averaging 1 mm. to 4 mm. in length are set in an aphanitic 
groundmass. 


Microscopically, many of these silicic volcanics are tuffaceous with fragmented 
phenocrysts in a once glassy mesostasis. Primary phenocrysts are quartz, 
plagioclase, alkali felspar, biotite, hornblende and rare pyroxene. Accessories 
include opaque oxides, sulphides, zircon and apatite. 


Quartz phenocrysts up to 4mm. across constitute 10% to 30% of the rock 
and typically show embayments. EHuhedral outlines suggest an original phase. 
Strings of tiny particles and inclusions of felspar, zircon and apatite are common. 
Fracturing of phenocrysts and undulatory extinction are characteristic. 


Plagioclase (10°% to 30% of the rock) occurs as subhedral laths and angular 
fragments, both compositionally identical. Hxtensive alteration to carbonate, 
sericite, epidote, prehnite and zeolites often obscures original compositions. 
Where determinations are possible cores are andesine and rims albite. Small 
laths are albitic (An,_,) while larger grains show patchy albitization. Potassic 
felspar phenocrysts are not abundant but locally may reach 207% in some flows. 
Usually the potassic felspar is surrounded by a rim about 0-2 mm. wide of albite 
as indicated by staining. Alteration to sericite and carbonate is widespread. 


Biotite (0-10°%) occurs as phenocrysts and replacing hornblende. Laths 
may reach 2mm. long. Most show kinked cleavages and shredding parallel 
to the basal cleavage. Hornblende phenocrysts are rare and often represented 
by small relict patches in areas of alteration. Pleochroism from olive green 
to mid brown is sometimes evident. Rare hypersthene may have been present 
more abundantly before alteration. The growndmass is typically cryptocrystalline 
and exhibits excellent flow structures. Staining shows, where grain size is 
resolvable, a granophyric (or granoblastic, if devitrified) intergrowth of quartz, 
potassic felspar and albite. Devitrified glass shards, following flow lineations, 
may constitute 50°% of the mesostasis. Arcuate lenses of minutely crystalline 
quartz and more rarely alkali felspar appear to result from devitrification (e.g. 
33729). Here, lenticular areas consist of parallel rows of tiny zeolite needles. 
Replacement of shards by chlorite has also taken place. 


e 


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290 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


The classification of these silicic volcanics as toscanites and dacites with 
tuffaceous equivalents is based on observed modes now evident in the rocks. 
Contact metamorphism—Petrography : The Brayton volcanics exhibit the effects 
of contact metamorphism in four main ways: 

(1) Grain growth of the mesostasis obliterates all evidence of primary 
structures. A granoblastic intergrowth of quartz and alkali felspar 
with minor biotite replaces the original groundmass. 

Original mafic phenocrysts show progressive changes. Low grade 
alteration of pre-existing hornblende, pyroxene and biotite produced 
prehnite, sphene, chlorite, opaque grains and epidote prior to contact 
metamorphism. Approach to the granitic contact shows development 
of green “‘ biotite ’’ (interlayered chlorite and mica ?), then hornblende, 
or brown biotite. 

Contact metamorphism of minerals in veins previously formed by low 
grade alteration, produced assemblages such as epidote-diopside-garnet- 
amphibole-quartz-albite. 

(4) Annealing recrystallization of large strained embayed quartz grains is 

evident (see Plate xiv, fig. 3). 

Three broad zones which represent an increase in metamorphic response 
are observable progressing towards the adamellite contact. Their recognition 
is based on the degree of recrystallization of the groundmass and mineralogical 
changes. 

33635 is typical of silicic volcanics on the outer edge of the aureole. The 
eroundmass shows moderate recrystallization with no remaining flow structures. 
No zeolite phases persist and tiny flakes of pale green amphibole are scattered 
throughout. Quartz phenocrysts are unaffected, alkali felspars are extensively 
altered to sericitic patches, while plagioclase is mostly albitic and clouded with 
epidote, carbonate, chlorite and _ sericite. Original mafic phenocrysts are 
completely pseudomorphed by epidote associated with cloudy aggregates of 
sphene and leucoxene, or interlocking felts or pale green mica with flakes of blue 
green amphibole and minor epidote. 

33648 shows a successive metamorphic response. The groundmass grain 
size has increased but quartz is still unaffected. Potassic felspar is completely 
altered to sericite and tiny quartz blebs. Original phenocrysts are all rimmed by 
a narrow zone, 0-1 mm. wide, of clear albite granules. Green-brown biotite and 
subordinate green chlorite occur. Outlines of original euhedral phenocrysts 
are no longer sharply defined. 

The highest metamorphic grade attained is shown in 33740 and 33755, which 
exhibit hornblende-rich and biotite-rich assemblages respectively. The ground- 
mass is granoblastic and the assemblages indicate probable attainment of upper 
hornblende hornfels facies conditions. Potassic felspar grains are not extensively 
altered and both felspars show incipient recrystallization. Quartz grains (see 
Plate xvi, fig. 3) show annealing crystallization where grains were originally 
Strained. 33740 contains clots and strings of hornblende laths up to 0-1 mm. 
long. These are pleochroic with X—deep blue green, Y=dark olive green 
and Z—=pale yellow green. 2V:x is ca. 70°, ZAc is ca. 20°. Associated sphene 
granules abound. 33755 exhibits decussate aggregates of biotite (Plate XVI, 
fig. 4) with each lath up to 0-5mm. long. Rare amphibole laths and opaque 
grains may be associated with biotite, while sphene is absent. Opaque grains, 
zircon and apatite are constant accessories. 

Thin sections 33645 to 33652 are representative of the explosive breccia 
discussed above. This outcrops at 29587139, close to the adamellite contact. 
Silicic voleanic fragments show zonation with an inner core of normal dacite or 
toscanite surrounded by a quartzofelspathic outer rim up to 15mm. wide 


(2 


— 


(3 


— 


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SUZANNE Y. O’REILLY 291 


comprising a zone leached of mafic constituents. These are set in a “ ground- 
mass’ of granular intergrowths of diopside, hedenbergite and epidote which 
may be penetrated by laths of amphibole. Irregular, clear quartz blebs, carbonate 
patches and garnet grains are evident. Rare porphyroblasts of albite occur, 
while tiny opaque and sphene granules may be locally abundant. These con- 
trasted compositions arise from an original chemically homogeneous rock as 
indicated by the migration of mafic components from fragment peripheries. 
While the inner core of fragments is compositionally equivalent to the normal 
silicic volcanics, their leached rims are higher in alkali and silica content, and the 
inter-fragment area is enriched in iron, calcium and magnesium. 


It has been suggested above that this breccia formed by explosive volcanic 
activity. Devitrification at low temperatures and pressures prior to granitic 
intrusions initiated chemical reorganization mainly involving the ‘‘ groundmass ”’ 
but also affecting fragment rims. The granite subsequently produced the 
observed assemblages (cf. Vallance, 1967). 


Metamorphism and previous concepts: The relationship of the granitic rocks to 
the silicic volcanics closely parallels that of the granodiorite and ‘‘ porphyrites ”’ 
south of Marulan. They were studied in detail by Osborne and Lovering (1952) 
and Svenson (1950), following Woolnough’s (1909) general observations. 


Osborne and Lovering (1952) considered the ‘‘ porphyrites ”’ were intruded 
as a high-level sill into Lower Palaeozoic country rocks. With the ‘‘ porphyrites ”’ 
still at elevated temperatures, intrusion of the granodiorite took place. Through 
slow cooling hybridization by interaction of magma and porphyrite ensued. 
Three ‘‘ hybrid zones ”’ were developed exhibiting mutual gradations representing 
successive degrees of hybridization. This process involved migration of soda 
from the magma to be deposited as albite rims in the hybrids. Extensive 
hydrothermai activity supplied ‘‘ vigorous solutions ’’ which effected formation 
of epidote, sericite and chlorite in hybrid ‘‘ porphyrites”’. Diffusion into 
hybrids of TiO, and alkalis as well as concentration of CaO and MgO to produce 
hornblende and biotite was also invoked. Analyses of one porphyrite, two 
hybrids and the granodiorite were recorded. 


No evidence of such chemical interaction was detected between rocks of 
the Brayton volcanics and the Lockyersleigh adamellite. A similar relationship 
was noted by Gould (1966) and Wass and Gould (1968) between the Tangerang 
voleanies and Glenrock granodiorite at South Marulan. 


The silicic voleanics show both extensive alteration and response to contact 
metamorphism. Observed characters may be attributed to a metamorphic 
progression towards the granite contact in rocks already altered rather than to 
development of ‘‘ hybrid zones’. These zones are probably correlative with the 
three stages of metamorphism discussed above. Compositional variations 
may be due to redistribution of components during previous low grade alteration 
and intrinsic chemical differences in individual flows. 


The sodic rims on alkali felspars do not necessitate addition of sodium. 
This phenomenon occurs even in rocks removed from the influence of the granitic 
rocks and may represent complete subsolvus unmixing of sodic and potassic 
phases. Formation of hornblende and biotite clots near the granitic contact 
does not require concentration of CaO and MgO as the chlorite and epidote of 
outer zones could contribute required components. 


Thus the present study of the Brayton volcanics indicates that a metamorphic 
progression rather than development of hybrid zones accounts for textures and 
mineral assemblages observed. 


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292 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


IGNEOUS INTRUSIONS 


Two components of the Marulan batholithic complex are represented. These 
are the Lockyersleigh adamellite and Towrang granodiorite. A contact meta- 
morphic aureole up to one mile in width is developed. Metamorphic assemblages 
produced in the intruded rocks have been discussed within each rock unit. 


Lockyersleigh Adamellite 


The Lockyersleigh adamellite was mapped over approximately three square 
miles (Fig. 1), but extends considerably southwards. Where exposed, contacts 
with country rock are sharp. Silicic volcanics of the Brayton volcanics were 
intruded by this adamellite and are cut by leucocratic veins derived from the 
intrusion. In places, exposures must be close to the roof of the intrusion as 
fine-grained phases are evident. Aplitic dykes, representing late-stage crystal- 
lization of the adamellite, contain miarolitic cavities lined with alkali felspar, 
smoky quartz and tourmaline crystals. Veins of epidote up to 2mm. wide 
occur throughout the adamellite. 


Petrography: The adamellite is holocrystalline, hypidiomorphic granular and 
shows a porphyritic texture with phenocrysts of alkali felspar, plagioclase, quartz 
and, rarely, biotite or hornblende. These minerals are duplicated in the ground- 
mass. Abundant sphene, opaque grains, epidote, zircon, apatite and alteration 
products including chlorite, carbonate, and sheaves of muscovite and prehnite 
are present. Quartz generally constitutes 30° to 40% of the rock. Phenocrysts 
are up to 4mm. across, anhedral and often embayed. Undulose extinction, 
Bohm striae and development of strain-free grains are common. Potassium 
felspar constitutes about 30% of the rock. Subhedral phenocrysts range from 
4mm. to 10mm. long. The optic axial plane is perpendicular to (010); 2Vx 
is approximately 50°; extinction on (001) is parallel to (010) cleavage, and on 
(010) ranges from 7 °” to 9°. Thus the felspar is monoclinic or has very low 
triclinicity. The latter is doubtful as ‘‘ tartan’ twinning is absent (Laves, 
1950). The composition lies in the range Or,, ..Ab+An,; 3, (Tuttle, 1952), 
and thus the felspar is an orthoclase low-albite perthite. Film and vein perthites 
are well developed: myrmekitic intergrowths are common. Plagioclase forms 
about 30°%% of the rock. Phenocrysts (2mm. to 4mm. long) show consistent 
core/rim compositional ranges of about An;, to about An,, respectively. 
Hornblende (absent in the Towrang granodiorite) occurs sporadically as euhedra 
or ragged subhedra; a reaction relationship giving biotite is often observed. 
It is pleochroic X=pale yellow, Y—grass green, Z—olive green. 2Vx is approxi- 
mately 70°. Biotite comprises up to 5°%% of the rock. Pleochroism is X =orange 
brown, Y =pale straw yellow, Z=deep speckled chocolate brown. 2V xis approxi- 
mately 10°. Alteration to chlorite, muscovite, prehnite, carbonate and granular 
epidote is common. Modal sphene may reach 5%. It occurs as granular 
aggregates, but commonly as euhedra 0-5 mm. to 2mm. long. Sphene is more 
abundant where hornblende is locally in excess of biotite. The intergranular 
groundmass forms 20% to 40% of the rock and comprises all mineral phases 
present in the rock. 


Towrang Granodiorite 

This granodiorite forms a small stock exposed at 294712. In outcrop it is 
approximately elliptical with east-west elongation. About one mile west of the 
area and one mile north of Towrang, a similar granodiorite is evident. These 
two masses may represent a single intrusion. 

The relationship with the Lockyersleigh adamellite is obscure because of 
poor outcrop on the boundary. The grey and white rock is quite uniform in 
hand specimen although occasional pegmatite patches are noted. These occur 


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SUZANNE Y. O’REILLY 293 


as veneer-like pods up to one foot in diameter, but never exceeding one inch in 
thickness. Distinct from the Lockyersleigh adamellite, no tourmaline occurs 
in the granodiorite but pyrite is common in the pegmatite phases. 


Petrography : In thin section the rock is holocrystalline, allotriomorphic granular. 
Minerals present are quartz, plagioclase, potassium felspar, biotite, opaque orains 
and accessory epidote, sphene, apatite, zircon, carbonate and muscovite. Quartz 
constitutes 30% to 40% of the mode and is typically anhedral and interstitia] 
although grains may obtain a maximum diameter of 3 mm. Well-developed 
Bohm lamellae and mosaic textures are evident. Plagioclase occurs as poorly 
terminated subhedra from 1mm. to 4mm. long and constitutes up to 40°% of 
the mode. Compositions range from an average of An,, for the most calcic cores 
to An, for outermost rims. Broad antiperthitic veins show rare development. 
Some large plagioclase grains have highly resorbed cores which appear to be 
accumulophyric. Vance (1962) attributes such textures to “ patchy zoning is 
arising from an intense phase of uneven resorption and subsequent re-precipitation 
and infilling by more sodic plagioclase. In rare grains with resorbed cores mosaic 
domains in optical continuity produce a patchiness on a 0-1 mm. scale and appear 
to control or be controlled by twin lamellae. 


Potassium felspar constitutes 20° to 30% of the rock and occurs as anhedra 
up to 4mm. across. The optic axial plane is perpendicular to (010) and straight 
extinction of (001) indicates a monoclinic structure. Extinction on (010) of 
10° to 12° indicates a composition Or,,Ab-++An,, to Or,,Ab+An,,. Perthitic 
textures are variably developed: albitic rims and large segregations are rare 
while no myrmekite was detected. 


Biotite tends to occur in aggregates and represents no more than 5° of the 
mode. Laths (0-2 to 1:0mm. long) are pleochroic with X—=straw yellow, 
Y=red-brown, Z—chocolate brown. Inclusions of zircon and apatite euhedra 
are common. 


Emplacement and Age 


Employing the genetic classification of Tuttle and Bowen (1958), the 
Lockyersleigh adamellite and the Towrang granodiorite both represent subsolvus 
or group II granites. All features observed are characteristic of plutons emplaced 
in epizonal regions (Buddington, 1959). 


The age of these intrusions can only be determined broadly as post-Brayton 
volcanics (Upper Silurian to Lower Devonian) and pre-Tertiary. Regarding the 
Marulan batholithic complex as a whole, Naylor (1939) records Upper Devonian 
sediments with a typical Lambiai fauna, resting unconformably on granitic 
rocks near Taralga and Bungonia. At South Marulan quarry, Upper Silurian 
limestone is metamorphosed by granite. Thus an age of Lower to Middle 
Devonian may be inferred on stratigraphic evidence. However, Evernden and 
Richards (1962), using the K/Ar ratio of constituent biotite of a granodiorite 
from Bungonia, determined an age of 307Xx10°® years (Carboniferous). This 
apparent anomaly may be due to non-contemporaneity of all phases comprising 
the Marulan batholithic complex or to loss of original argon. 


TERTIARY ROCKS 


Two basaltic flows (see Fig. 1), the Billyrambija dolerite mass and scattered 
sediments represent Tertiary rocks present. Age of the igneous rocks is inferred 
from their intimate association with sediments containing Tertiary flora and 
from their petrological affinity to basaltic rocks in the Moss Vale region which 
have a Tertiary age as shown by potassium-argon dating (P. Wellman, pers. 
comm.). 


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294 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


Tertiary Sediments 


Isolated small outcrops of Tertiary sediment are associated with the 
Billyrambija dolerite where they occur mainly at the base and the 100 feet contour 
level, and with the two flows. Lamination of relatively fine and coarse fractions 
on a centimetre scale is marked, each layer often exhibiting graded bedding. 
Detritus comprises dominant volcanic, plutonic and reworked quartz grains, 
together with altered felspar grains and aggregates, while hornblende, epidote 
and muscovite are accessories. This detritus appears to be derived directly 
from nearby sources such as the Brayton volcanics, the silicic intrusions and the 
Ordovician or (?) Silurian sediments. Extreme angularity of grains suggests 
little transportation. The interstitial material may be partly matrix and partly 
cement and constitutes from 20° to 30% of the rock. It consists of very fine- 
grained, heavily iron-stained siliceous material. 

The flora is poorly preserved, but most specimens are dicotyledonous leaves 
belonging to the Cinnamomum flora. One well-preserved fragment is similar to 
Cinnamomum polymorphoides, McCoy, 1874. That the leaves may be placed in 
the family Lawraceae and compared with species of Cinnamomum is indicated by 
comparison with similar but well-preserved fossil leaves from Wingello, which 
are considered to belong to the Cinnamomum flora (D. Selkirk, pers. comm.). 
This flora is Tertiary or, less probably, late Cretaceous in age. However, such 
leaves are termed Tertiary as geologists, on topographic and physiographic 
evidence, have considered closely associated basalts to be Tertiary. The gradual 
accumulation of potassium- -argon dating of such basalts (e.g. McDougall and 
Wilkinson, 1967) supports this in general and thus, with available data, it seems 
reasonable to consider these sediments Tertiary. 


Billyrambya Dolerite 


The Billyrambija dolerite has a maximum exposed height of 340+10 feet 
and outcrops in an approximately circular pattern with a diameter of one mile. 
Field relations do not provide conclusive evidence on the mode of emplacement. 
However, it is suggested that it may represent a very high level intrusion, 
possibly disrupting unconsolidated sediment and reaching the surface in places. 
This is supported by the presence of chilled phases which form randomly placed 
knolls and shoulders, and the presence of blocks of Tertiary sediment at the 
100 feet contour line which show random bedding orientations. 


Three components of the dolerite are recognized : chilled phases, dykes and 
the main dolerite body. The chilled phases occur associated with Tertiary 
sediment and are mineralogically comparable to the dolerite. Phenocrysts of 
titanaugite rarely exceed 2 mm. in diameter ; olivines are typically skeletal and 
clouded with inclusions, are unzoned and have a composition about Fo,, 
(8=1-680-40-003). Plagioclase (Ang core to An,, margin) is abundant and 
laths often show swallow-tail terminations. The groundmass is extremely 
fine-grained but tiny pyroxene and opaque granules are evident. Dyke rocks 
are distinctive in mineralogy and field occurrence. Dykes apparently intruding 
the dolerite can usually be traced over a vertical height of 20 feet and vary 
from five feet to one foot in width. Wider dykes exhibit chilled margins. 
Euhedral titanaugite, olivine and plagioclase (core An,,, margin An, )) are set 
in a fine-grained groundmass dominantly composed of aegirine-augite and 
magnetite. Pyroxene phenocrysts may be mantled by aegirine-augite and are 
often densely clouded with magnetite granules. Olivine phenocrysts are zoned 
from Fo,, to Fo,, (core and margin respectively). Amygdales abound in dyke 
rocks and are filled with thomsonite, chabazite, natrolite, analcite, prehnite or 
clay minerals. 

The Billyrambija dolerite comprises pyroxene (35-40%), olivine (13-28%), 
plagioclase (14-25°%) and opaque grains (3%) set in a mesostasis (14-25°%) of 


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SUZANNE Y. O'REILLY 295 


zeolites, tiny opaque granules, laths of K-felspar, flakes of aegirine and 
(2) kaersutite, analcite and secondary chlorite. Apatite needles up to 0-5 mm 
long abound. 

Titanaugite euhedra occur in two generations. Those of the first generation 
reach 5mm. across, may occur singly or in glomeroporphyritic aggregates, and 
are commonly sub- “ophitically indented by plagioclase. Inclusions of a cubic 
iron-titanium oxide, olivine and a colourless (?) pyroxene are common. Pleo- 
chroism with X =pinkish yellow, Y =pink, Z=mauve-pink is variable in intensity. 
Well-developed hour-glass structures are present. Zoning is striking and 
generally limited to the outer few millimetres of the large euhedra. Second 
generation pyroxenes are prismatic and reach a length of 0-5 mm., lack significant 
zoning, and are optically identical to the outer margins of large pyroxenes. 


Olivine also occurs in two distinct generations, as subhedral to anhedral 
grains up to 0-5 mm. across and as euhedra up to 2mm. long. The latter have 
compositions from about Fo,, to Fo,. as determined by refractive index and X-ray 
methods. 

Plagioclase occurs as euhedral, zoned laths 0-5-2-0 mm. long. Cores exhibit 
compositional variation with upward progression from An,; to cae rims being 
up to 15 mol. per cent. richer in Ab. 


Opaque minerals identified were ilmenite and titanomagnetite. Ilmenite 
occurs as laths which may be sub-dendritic in habit and represent 10° of opaque 
minerals present. Titanomagnetite occurs as octahedra up to 0-:2mm. wide. 
Compositions were estimated by measuring cell edges and using the curve of 
Basta (1957). Titanomagnetite from the lowest level was approximately 96% 
Fe,TiO,, that from the highest level 70% Fe,TiO,. 

Modal analyses indicate that mild differentiation may have taken place 
within the dolerite body: there is an upward decrease in mode of olivine and 
plagioclase and increase in volume percentage of mesostasis. This is supported 
by the limited and preliminary (crop O stun data on the plagioclases and 
titanomagnetites. 


Tertiary Basaltic Flows 


Two tertiary basaltic flows are evident (Fig. 1). The southerly flow is 
petrographically identical with the chilled phase of the Billyrambija dolerite. 
Perhaps the original extent of this latter body was greater than now observed, 
this flow representing a southerly projection of the Billyrambija dolerite. 

The northerly flow is petrographically distinct, although still a typical 
alkali basalt. Extensive alteration, mainly due to weathering, has taken place. 
Pyroxene is rare ; olivine euhedra up to 1-0 mm. long comprise 10°% of the rock 
and are extensively altered. Plagioclase laths (30°% of the rock) are zoned from 
An, (core) to An,,; (margin) and define a flow lineation. The groundmass may 
have been glassy ; felspar microlites abound as well as fine opaque particles. 


ACKNOWLEDGEMENTS 


I wish to thank Associate Professor T. G. Vallance, who supervised the 
project, and who with Dr. R. H. Vernon critically read the manuscript. 
Laboratory facilities at the University of Sydney were made available by 
Professor C. E. Marshall. 


References 
Basta, E. Z., 1957.—Accurate determination of the cell dimensions of magnetite. Mzneralog. 
Mag., 31: 431-442. 
Browne, W. R., 1933.—An account of post-Paleozoic igneous activity in New South Wales. 
J. Proc. R. Soc. N.S.W., 67: 9-95. 
Buppineton, A. F., 1959.—Granite emplacement with special reference to North America. 
Bull. geol. Soc. Amer., 70: 671-747. 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, VoL. 96, Part 4 


296 PETROLOGY AND STRATIGRAPHY, BRAYTON DISTRICT, N.S.W. 


CLARKE, W. B., 1845.—On dykes of marble and quartz in connection with plutonic rock on the 
Upper Wollondilly in Argyle County, New South Wales. Q. Jl. geol. Soc. Lond., 1: 342-344. 

CraFt, F. A., 1928.—Physiography of the Wollondilly River Basin. Proc. Linn. Soc. N.S.W., 
53: 618-650. 

EVERNDEN, J. F., and Ricuarps, J. R., 1962.—Potassium-Argon ages in eastern Australia. 
J. geol. Soc. Aust., 9: 1—50. 

GouLp, I. G., 1966.—The general geology of an area south of Marulan. Unpubl. hons. thesis, 
University of Sydney. 

LavEs, F., 1950.—The lattice and twinning of microcline and other potash feldspars. J. geol. | 
58: 548-571. 

Loverinec, J. F., 1950.—Contribution to a study of the Marulan Batholith. Unpubl. hons. 
thesis, University of Sydney. 

McDoveatt, I., and Witxrinson, J. F. G., 1967.—Potassium-Argon dates on some Cainozoic 
voleanic rocks from north-eastern New South Wales. J. geol. Soc. Aust., 14: 225-233. 
Naytor, G. F. K., 1935a.—Note on the geology of the Goulburn district with special reference to 

Palaeozoic stratigraphy. J. Proc. R. Soc. N.S.W., 69: 75-85. 
, 1935b.—The Palaeozoic sediments near Bungonia ; their field relations and graptolite 
fauna. Tbid., 69: 123-134. 
, 1938.—Graptolites of the Goulburn district. Jbid., 72: 129-135. 
, 1939.—The age of the Marulan Batholith. Jbid., 73: 82-85. 
, 1949. A further contribution to the geology of the Goulburn district. Jbid., 83: 
279-287. 
OsBorRNE, G. D., 1948.—The contaminated granodiorites of South Marulan and Marulan Creek, 
Part 1. Jbid., 82: 116-128. ; 
, and LovERING, J. F., 1952.—Contributions to a study of the Marulan Batholith. Ibid. 
86: 108-118. 
Svenson, D., 1950.—Contribution to a study of the Marulan district. Unpubl. hons. thesis, 
University of Sydney. 
Tuomas, D. E., 1960.—The zonal distribution of Australian graptolites. J. Proc. R. Soc. N.S.W.., 
94: 1-58. 
TuTTLeE, O. F., 1952.—Optical studies on alkali felspars. Am. J. Sci... Bowen Volume: 553-567. 
, and Bowen, N. L., 1957.—Origin of granite in the light of experimental studies in the 
system NaAIS8i03;-KAI8i03-Si0,-H,O0. Mem. geol. Soc. Amer., 74. 
VALLANCE, T. G., 1960.—Concerning spilites. Proc. Linn. Soc. N.S.W., 85: 8-52. 
, 1967.—Mafic rock alteration and isochemical development of some cordierite-antho- 
phyllite rocks. J. Petrology, 8: 84-96. 
VANCE, J. A., 1962.—Zoning in igneous Plagioclase-normal and oscillatory zoning. Am. J. Scv., 
260: 746-760. 
Wass, R. E., and Goutp, I. G., 1968.—Permian faunas and sediments from the South Marulan 
district, New South Wales. Proc. Linn. Soc. N.S.W., 93: 212-226. 
WootnoucH, W. G., 1909.—The general geology of Marulan and Tallong, New South Wales. 
Ibid., 34: 782-808. 


PROCEEDINGS OF THE LINNEAN SOCIETY OF NEw SourH WALES, VoL. 96, Part 4 


Proc. LINN. Soc. N.S.W., Vol. 96, Part 4 PLATE XVI 


EXPLANATION OF PLATE XVI 


1. Flow foliation in an altered dacite outside the contact aureole, 33729. x 45. 

2. Devitrified glass shards in a dacitie tuff outside the contact aureole, 33734. x 45. 

3. Embayed quartz phenocryst showing incipient recrystallization in a dacite within the contact 
aureole, 33728. x45. 

4. Biotite clot and hornfelsic texture in a metamorphosed acid voleanie rock adjacent to the 
adamellite contact, 33655. 45. 


SEDIMENTATION IN JERVIS BAY 


ee Gay AanOR: 
Department of Geology, Australian National University 


(Communicated by Dr. K. A. W. Crook) 
[Accepted for publication 23rd June 1971] 


Synopsis 
Jervis Bay is underlain by a lenticular sand body occupying a river valley cut in Permian 
sandstones and siltstones. The sediment covers an area of 84km.? with an average thickness 
of 20m. 
Two zones are evident, a low energy central zone consisting of a sublitharenite containing 
less than 85% quartz and a marginal high energy zone containing quartz arenites and sub- 
litharenites containing greater than 85% quartz. 


SAMPLE LOCALITY | 


MAP 


SYDNEY 


JERVIS 
BAY 

360s 

152°E 


Huskisson 


“85 "87 +89 “9 “93 Beecratt 


Vincentia Bay 
2206 +74 78 +80 
Plantation Pt. 


Peninsula 


Vincentia 


-42 744 746 S 
Beach “47 5 Tasman Sea 
Sf 
+26 -28 0) oil 233 2a 
: Bi i= 
Sailors ra 
18 
6 . Bowen 
Pe Is: 
= 
Darling Road s 
we Gs  % 
fc 
60 
= 
or 
> 
) 
O 0 1 2 
Bherewerre — SS ee 


Peninsula Kilometres 


Fig. 1. Map showing local names referred to in the text and sample localities. 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH WaAtgES, VoL. 96, Part 4 


SEDIMENTATION IN JERVIS BAY 


bo 
ite) 
(06) 


INTRODUCTION 


The recent marine sediments in Jervis Bay have been investigated (Taylor, 
1970) and an attempt made to delineate the character of the sediments, the 
processes involved in sedimentation, and the history of the embayment. This 
was carried out by surface sampling (Fig. 1), continuous seismic profiling and a 
detailed examination of the hydrology, bathymetry and Quarternary geology 
of the bay. . 

REGIONAL GEOLOGY 

Jervis Bay is an embayment in the coastline of New South Wales approxi- 
mately 84 km.? in area with its entrance 6 km. east of the general line of the 
coast. Itis situated in a gently folded sequence of Lower Permian sublitharenites 
and siltstones (Rose, 1966) (Fig. 2). Permian rocks crop out over half of the 
coastline of the bay; the remainder is bordered by beaches and beach ridges 
dating back to the Pleistocene (Walker, 1967). 


Only one stream of any significance, Currambeen Creek, drains into the bay, 
and most of its sediment load is deposited before reaching the bay. Other 
streams deposit their load in back-swamps and lagoons around the bay. 


WS" é a GEOLOGY 


A Lh Cn 
—— 


Kilometres 


LEGEND 
[o] ca 
VOLCANICS? —~—” CATCHMENT BOUNDARY 
7 — 

NSN opeRRY SHALE _®” ANTICLINE 
Z NOWRA SANDSTONE eS SYNCLINE 
= Ff 
ce = WANDRAWAN DIAN r-7 FAULT 
= SILTSTONE 

5 

CONJOLA FORMATION — OoDIP 


NO SURFICIAL GEOLOGY SHOWN 


Fig. 2. Catchment Geology of Jervis Bay and of streams entering the bay. 


PROCEEDINGS OF THE LINNEAN SOcIETY oF NEW SoutH WALES, Vou. 96, Part 4 


G. TAYLOR 299 


BATHYMETRY 
R.A.N. Chart AUS. 80 shows that the bottom slopes gently to 26 m. from the 
north, west and south, the eastern shoreline sloping more steeply (Fig. 3). The 
entrance to the bay reaches a maximum depth of 41 m. 


BATHYMETRY 


8 cng ee FROM R.A.N. CHART AUS. 80 
12 


Kilometres 


CONTOURS IN METRES 


Fig. 3. Bathymetry of Jervis Bay. 


HYDROLOGY 


No full-scale hydrological survey has been conducted in Jervis Bay. <A 
limited survey was conducted in the southern half of the bay by the author in 
the summer of 1968. Three water layers were present: an upper layer up to 
17 m. thick, with an intermediate layer up to 4:5 m. thick, and a lower layer up 
to 20 m. thick in the deeper parts of the bay. The mean summer surface and 
bottom temperatures are 23-1°C. and 19-8°C. respectively and average 
chlorinites are 19-65%,. The bay has a similar hydrographic environment to 
other open estuaries in eastern New South Wales (Rochford, 1951). 


PROCEEDINGS OF THE LINNEAN Society or New SourH WatgEs, Vou. 96, Part 4 


300 SEDIMENTATION IN JERVIS BAY 


The predominant swell entering the bay is from the south-east, causing 
turbulence on exposed shores all year to a depth of 19m. Predominant winds 
are north-easterly and north-westerly, with southerly storm winds. Hence 
all shorelines within the bay are subject to continual wave activity, except 
south-facing shores at the north end of the bay, which are only affected during 
southerly storms. 


SORTING 


J 
Sie 
“5 SC 
] 
0): -0 ] 2 
cole. ys 
Kilometres 


CONTOURS IN ® 
UNITS 


Kilometres 


Fig. 4. Maps showing the surface distribution of grain size, sorting and sand/mud ratios in 
Jervis Bay. 
HIELD AND LABORATORY PROCEDURES 


Grab samples of the top 15 cm. of sediment were taken at 1-6 km. intervals 
on an H.-W. grid and at 0:8km. on a N.-S. grid. The samples were split, 
desalinated, and dried in preparation for mechanical and mineralogical analysis. 


PROCEEDINGS OF THE LINNEAN SociETY oF NEw SoutH WALES, Vou. 96, Part 4 


G. TAYLOR 301 


MECHANICAL ANALYSIS 


Measurements of grainsize, sorting and sand/mud ratios were made using 
techniques described by Folk (1968). The mean size and sorting coefficients 
were calculated from frequency data using formulae derived by Folk and Ward 


(1957). 


The mean size of sediments, excluding the mud fraction, varies from 0-649 


to 3-379. 


The areas of coarse sediment are confined to regions of high energy 


in the entrance to the bay. The general trend in the bay is a fining marginwards 
(Fig. 4); off Callala Beach and Darling Road this trend is masked by growth of 


FELDSPAR 


ROCK 
FRAGMENTS 


te) 1 2 


Kilometres Kilometres 


PETROGRAPHIC 
PROVINCES 


Quartz Arenite 
Sublithic 


Arenite 


95 a Quartz Arenite 
/ kX Sublithic 


Arenite 


@)y Ul te 
_——— 
Kilometres 


Kilometres FELDSPAR ROCK FRAGMENTS 


Fig. 5. Maps showing the surface distribution of quartz, rock fragments, feldspar and 
petrographic provinces in Jervis Bay. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH Waters, Vou. 96, Part 4 


302 SEDIMENTATION IN JERVIS BAY 


molluscan fauna. The mud fraction has a mean size in the coarse silt range 
(4:76 to 5-01) with less than 5% being finer than 79. 


The marginal sediments are slightly better sorted than those towards the 
centre (Fig. 4). The coarse sediments in the entrance and in areas of molluscan 
growth are poorly sorted. The sorting values vary from 1-909 units to 0-339 
units, and the areal distribution pattern follows that for mean size, i.e. the 
better sorted sediments occur in marginal regions and more poorly sorted areas 
are central. 


No sediment contains more than 20°% mud, and most contain less than 1%. 
The sand/mud ratio (Fig. 4) varies from infinity to 1-75. The regions of high 
mud concentration occur in areas protected from the south-easterly swell, e.g. 
Vineentia Bay, Green Point and off South Sailors Beach. 


MINERALOGY 


The composition of the terrigenous sediments was determined by point 
counting, and the amount of carbonates by solution in hydrochloric acid. 


LIGHT MINERALS 


The abundance of quartz varies between 73°, and 96° of the total 
terrigenous constituents. The higher concentrations are restricted to the marginal 
regions (Fig. 5). The majority of quartz is well rounded and monocrystalline, 
indicating a high degree of maturity. 

Rock fragments make up between 3% and 21%, the highest concentrations 
being in the central region (Fig. 5). The majority of rock fragments are sandstone 
from the seacliffs surrounding the bay. Minor basic igneous and siltstone 
fragments are also found. These rock fragments are all unstable and reworking 
destroys them, hence their paucity in the marginal zones. 

Feldspar occurs in minor amounts in the central region (Fig. 5) and in areas 
of active erosion of the Nowra Sandstone (Rose, 1966). It is mostly highly 
altered and contains included mica. It is mostly a sodium-calcium variety with 
little potassium feldspar. 


CARBONATES 


The carbonates locally contribute up to 80°% of the sediment in the areas of 
strong tidal currents near the entrance, and of abundant faunal growth. The 
greater part of the sediments however contain between 5% and 20°% carbonate 
(Fig. 6). The carbonates are made up of complete and fragmented remains of 
bivalves, gastropods, echinoids, forams, polyzoans and ostracods, the last four 
being more common in the southern region of the bay. 


PETROGRAPHIC PROVINCES 


The petrographic sand types have been classified according to Folk < (1968). 
Sublitharenites cover most of the bay with small areas of quartzarenite at the 
north and south end (Fig. 5). 


The abundance of quartz in marginal zones of high energy and fine grain 
size and the comparative lack of rock fragments in these zones is presumably 
due in part to the break-up of rock fragments. Since the rock fragments are 
mainly of a sublitharenite, the contribution to the sediments from their break- 
down is essentially quartz. This process is effected less rapidly in areas of lower 
energy (i.e. central region) and hence rock fragments survive longer. 


The erosion of Nowra Sandstone (Rose, 1966) around the eastern margin 
of the bay continuously supplies rock fragments in that region, hence although it 
is a high energy coast, rock fragments are relatively abundant. 


PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SoutH WALES, VoL. 96, Part 4 


G. TAYLOR 303 


Feldspars are more abundant in the deeper lower energy zones of the bay, 
presumably because of their instability in areas of continual wave activity. 

These provinces are masked to some extent by abundant carbonate detritus 
present in some localities. 


i, CARBONATE 


Kilometres 


Fig. 6. Map showing the surface distribution of the carbonate components of the sediment. 


HEAVY MINERALS 

The heavy minerals from 10 g. splits of sediment were separated by use of 
tetrabromethane. The sizes of the grains separated varied from 0-08 to 0:12 mm. 
and from less than 0-01°% to 0:9°% of the marine samples by weight and up to 63% 
in one beach concentrate. The separates were then examined and ‘“ species ”’ 
counts made on at least 200 grains. Highteen different ‘“‘ species ’’ were noted, 
of which 50°% (mean percentage) were opaques, including magnetite, hematite, 
leucoxene, pyrite, cassiterite and ilmenite. Minor constituents (monazite, 
rutile, hornblende, andalusite, corundum, garnet, spinels and biotite) make up 
14%. The remaning minerals, zircon (2:5°%), tourmaline (18%) and epidote 
(16%), are of environmental significance. 

The provenance of these minerals is the Permian sediments in the catchment 
and seacliffs and from reworking of older unconsolidated sediments, themselves 
originating essentially from the Permian. 


From the heavy mineral provinces (Fig. 7) 


(zircon province Ais 8% zircon 
Tourmaline province 20% tourmaline 
epidote province ont ty LDA . ) 


it is clear that zircon predominantly occurs around the margins of the bay and 
that tourmaline and epidote are restricted to the deeper central areas. 

These distributions reflect the energy conditions in the bay and the specific 
gravity of zircon and tourmaline/epidote. .The heavier mineral, zircon, is 
concentrated in the high energy zones and the lighter minerals, tourmaline and 
epidote, are concentrated in the deeper low energy zones (Fig. 7). 


PROCEEDINGS OF THE LINNEAN Society oF New SoutH Wates, Vou. 96, Part 4 


304 SEDIMENTATION IN JERVIS BAY 


Mup FRACTION MINERALOGY 
Between 40% and 60°% of the muds are composed of biogenic carbonate 
debris. A variety of terrigenous components are present in the muds, the most 
abundant being quartz (>90°% of the total terrigenous fraction in all samples 
analysed). Other minerals found in minor percentages only include feldspar, 
kaolinite, chlorite, heavy minerals and muscovite. 


SEDIMENTARY HISTORY 


A continuous seismic reflection survey of 10 traverses (Fig. 8) using a low 
output ‘“ sparker”’ system with a visual display recorder, showed that Jervis 
Bay is an infilled river valley formed during a Pleistocene low stand of sea level 
(Fig. 8). The sediments deposited in these valleys have since been reworked 
and redistributed by at least one marine transgression. Examination of the 
records shows some patches of relict sediment which have been interpreted as 
lag-deposits of either the original fluvial sediments or sediments redistributed 
from an earlier marine transgression. 


HEAVY MINERAL 


ZIRCON/ 
PROVINCES == ES] zircon 


TOURMALINE 


EN tourmating 


Sy. ED epipote 


Kilometres 


_———s 
Kilometres 


Fig. 7. Heavy mineral provinces and zircon/tourmaline ratios of the surface sediments in 
Jervis Bay. 


Another source of sediment is the erosion of the seacliffs during the various 
interglacial high sea levels. No sediment is entering the bay from the shelf 
under present conditions ; however, this may not have been so during Pleistocene 
regressions and transgressions. The contribution of sediment from outside the 
bay while the bay was open at Hare Bayisunknown. A.D. Albani (pers. comm.), 
from seismic profiles in the Hare Bay region, confirms that the bay was at one 
time open to the sea at the north end also. Sediment in the bay is up to 30 m. 
thick in the deeper central areas, thinning to zero at the margins and over a 
basement high in the entrance region (Fig. 8). The thickness under the areas of 
beach-ridge development (e.g. Callala Beach) is unknown. 

Evidence of sedimentary structures is absent from both the seismic records 
and from grab samples. This is probably due in part to disturbance of the 
materials during sampling; however, in the few undisturbed samples taken 
few structures were apparent, owing to destruction by bioturbation. 


PROCEEDINGS OF THE LINNEAN SociETY or New SoutH Wates, Vou. 96, Part 4 


G. TAYLOR 305 


7 /AREATION 
OF SEISMIC 


TRAVERSES 


Kilometres 


BASEMENT 
CONTOUR 


30 
o\ SEDIMENT 
O}SiSOPACH 
MAP 


el 
Kilometres 


Veer 


CONTOURS IN METRES 
Kilometres eg ISOPACHS IN METRES 


Tig. 8. Maps showing the location of seismic traverses, the configuration of the solid sub- 
stratum and an isopach map of the unconsolidated sediments. 


PROCEEDINGS OF THE LINNEAN Society or New SourH Wates, Vou. 96, Part 4 


306 SEDIMENTATION IN JERVIS BAY 


DISCUSSION 


Sedimentation in Jervis Bay forms two distinct zones, the centre and the 
margin. These zones of deposition are based on differences between sediments 
forming in them. 

The grain size in the marginal zone is finer than that in the centre (2-440 
and 2-11 respectively). This results from the more intense reworking by wave 
activity in the marginal zones and the removal of silt, clay and carbonates to the 
central zone. The carbonate grains are significantly different in shape from the 
bulk of the sediment in the marginal zone and are hence unstable in this area. 
Thus it follows that the central zone is enriched in silt, clay and carbonates 
with respect to the marginal zone. 


As a result of the continual reworking in the marginal zone and the rejection 
of unstable particles, marginal sediments are better sorted (0-490 units compared 
to 0-69 units) than those in the central zone. The energy difference between 
the central and marginal zones also controls the mineralogy. Sedimentary rock 
fragments are broken down into their constituents, the major one being quartz. 
Hence quartz is more abundant in the marginal zone than in the central zone 
and rock fragments are more abundant in the central zone. 


The heavy minerals also reflect the differences in energy distributions ; 
the heaviest (i.e. zircon) concentrates in the high energy zones, and tourmaline 
and epidote in the low energy zones. 


In summary, the sediment distribution is controlled almost entirely by 
energy regimes in the bay. The only effective variable with respect to this in 
the bay is wave activity. 


ACKNOWLEDGEMENTS 


This study was carried out for a Master’s degree at the University of New 
South Wales in part under a Commonwealth Postgraduate Scholarship. The 
author wishes to acknowledge help, during all stages of the work, from Dr. A. N. 
Carter of the University of New South Wales and Dr. K. A. W. Crook of the 
Australian National University. 


References 
Foux, R. L., 1968.—Petrology of Sedimentary Rocks. Hemphills, Austin, Texas. 
, and Warp, W. C., 1957.—Brazos River: A study in the significance of grain-size 
parameters. J. sedim. Petrol., 27: 3-36. 
Rocurorp, D. J., 1951.—Studies in Australian estuaries, Hydrology. Aust. J. Mar. and Fresh- 
water Res., 2: 1-116. 
Rosz, G., 1966.—Ulladulla 1-250,000 Geological Map. New South Wales Mines Department. 
Taytor, G., 1970.—Sedimentation in Jervis Bay. M.Sc. thesis, Univ. N.S.W., unpubl. 
Watxer, R. G., 1967.—The coastal geomorphology of the Jervis Bay Area. M.Sc. thesis, A.N.U. 
unpubl. 


PROCEEDINGS OF THE LINNEAN SociETY or NEw SoutH WALEs, VoL. 96, Part 4 


ABSTRACT OF PROCEEDINGS 


ORDINARY GENERAL MEETING 
dlst March 1971 


Mr. L. A. 8. Johnson, President, in the chair. 

Minutes of the last Ordinary General Meeting (25th November 1970) and 
of the Special General Meeting (25th November 1970) were taken as read and 
confirmed. 

Suggestions for 1971 Programme: 


The Chairman invited comments and suggestions from members for meetings 
during 1971. After discussion, it was resolved that these would be passed on to 
the incoming Programme Committee for consideration. 


It was also resolved to recommend that Council consider abolishing the 
printed abstract of Proceedings, and that Council investigate the possibility of 
circulating brief quarterly summaries of Council activities. 

Council subsequently resolved to abolish printing the Abstract and substitute 
a roneoed circular instead. 

Paper read (by title only, with an opportunity for discussion at the next 
Ordinary General Meeting) : 

The Trichomes of the Goodeniaceae. By R. C. Carolin. 


ORDINARY GENERAL MEETING 
28th April 1971 
Held in the Committee Room, Institution of Engineers, Science House. 
Mr. L. A. 8. Johnson, President, in the chair. 
Minutes of the last Ordinary Meeting (31st March 1971) were read and 
confirmed. 
Announcements : 


The Secretary announced that the First International Congress of Systematic 
and Evolutionary Biology would be held at the University of Colorado, Boulder, 
U.S.A., in August 1973. Details could be obtained from the Secretary on request. 


The President announced with great regret the death of Professor H. N. 
Barker on 16th April 1971. Professor Barber had been a Life Member of the 
Society since 1950. 

Papers read (by title only) : 


Australian sea stars of the genus Patiriella (Asteroidea, Astirinidae). By 
A. J. Dartnall. 


A sea star of genus Ctenodiscus from Tasmania. By A. J. Dartnall. 

Late Devonian conodonts from the Luton Formation, northern New South 
Wales. By G. M. Philip and J. H. Jackson. 
Paper taken as read at the March Ordinary General Meeting (with an opportunity 

fer discussion at the April Ordinary General Meeting) : 

The Trichomes of the Goodeniaceae. By R. C. Carolin. 
Papers read: 

1. The genus Nwunciella Roewer (Opiliones, Laniatores) with description 
of a new species from Kangaroo Island, South Australia. By G.S. Hunt. 

2. The reptiles of Lord Howe Island. By H. G. Cogger. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WateEs, Vou. 96, Part 4 


308 ABSTRACT OF PROCEEDINGS 


ORDINARY GENERAL MEETING 
30th June 1971 


Held in the Committee Rooms, Institution of Engineers, Science House, 
at 7.30 p.m. 
Dr. L. A. S. Johnson, President, in the chair. 


Announcements : 
The President announced the conditions adopted by the Council of the 
Royal Society of N.S.W. for the award of the Walter Burfitt Prize. 


The President read a statement by the Minister for the Department of the 
Vice-President of the Executive Council regarding the formation of the Scientific 
and Technological Information Services Enquiry Committee. 


The following meetings of the Royal Society of N.S.W. were announced : 

7th July, at 7.45. (An address on 150 years of scientific associations. The 
Royal Society is celebrating its 150th anniversary.) 

4th August, at 7.45. An address by Colonel Magee, Sydney Cove Re- 
development Authority. 
Papers read : 

A bacterial leaf spot of zinnia in New South Wales. By A. L. Bertus and 
A. C. Hayward. 


An undescribed species of Ustilago on Cyperus lucidus R.Br. in Australia 
with comments on Robert Brown’s type collection of C. lucidus. By J. Walker. 


Shoot blight of Hucalyptus spp. caused by an undescribed species of 
Ramularia. By J. Walker and A. L. Bertus. 
Address : 


An address was given by Dr. J. 8S. Beard, Director, Royal Botanic Gardens, 
on ‘‘ New light on some aspects of geomorphology in Western Australia afforded 
by vegetation mapping’. 


ORDINARY GENERAL MEETING 
28th July 1971 

Held in the Committee Rooms, Institution of Engineers, Science House, 
at 7.30 p.m. 

Dr. L. A. 8S. Johnson, President, in the chair. 

Announcements : 

The President announced that there would be a Field Day on 19th September 
1971. Members would be notified later of further details. This would replace 
the ordinary monthly meeting for September. 

Papers read: 

The following papers were read by title only: 

New Species of Lophopetalum (Celastraceae), a genus new for Australia. 
By N. Byrnes. 

The food and feeding of winter whiting (Sillago maculata Quoy and Gaimard) 
in Moreton Bay. By John L. Maclean. 

Paper read: 


The early history and relationship of the New Guinea highland dog (Canis 
hallstromi). By KE. LeG. Troughton. 


PROCEEDINGS OF THE LINNEAN SociETY or NEw SoutH WALES, Vou. 96, Part 4 


ABSTRACT OF PROCEEDINGS 309 


Address : 


An address was given by Mr. 8. Jacobs, Royal Botanic Gardens, Sydney, 
entitled ‘‘ New Approaches to Grass Systematics ”’. 


ORDINARY GENERAL MEETING 
19th September 1971 


A Field Day Excursion to Dharug National Park replaced the Ordinary 
General Meeting in September. 


ORDINARY GENERAL MEETING 
27th October 1971 


Held in the Committee Room of the Institution of Engineers, Science House, 
Sydney, on Wednesday, 27th October 1971, at 7.30 p.m. 


Dr. L. A. 8. Johnson, President, in the chair. 


Announcements : 


The President announced the establishment of the Keith Hindwood Memorial 
Fund. 

Although the form of a memorial would depend on the amount of money 
donated, it was hoped that a Bird Studies Centre might be established. This, 
if it developed along lines of similar institutes abroad, would be available to the 
amateur student, a place where he could have access to a scientific library, 
record material, and which would be a nerve centre to concentrate and stimulate 
work by amateurs. 

Should enough money be raised, it might be possible to buy headquarters 
and pay for modest staffing, with options for future growth, although voluntary 
staff could be used at the beginning. 

The Trustees of the Australian Museum had generously consented to set up 
this Memorial Fund, and this meant all donations would be tax-deductible, 
cheques to be made payable to the Australian Museum Keith Hindwood Memorial 
Fund. 


Papers read: 


Australasian Ceratopogonidae. Part XV. The genus Alluwaudomyia Kieffer 
in Australia and New Guinea. By M. L. Debenham. 


Papers read by title only: 

The reaction of seedlings of Crtrus spp. and related genera to Phytophthora 
citrophthora. By P. Broadbent, L. R. Fraser and Y. Waterworth. 

Systematic position of the genera Triodia R.Br. and Plectrachne Henr. 
(Gramineae). By 8S. Jacobs. 

Aphids of Papua and New Guinea: An annotated check list. By K. P. 
Lamb. 

The phenology of species common to three semi-arid grasslands. By 
O. B. Williams. 


Symposium : 

A symposium was given by Mr. G. P. Whitley, Mr. G. N. Baur and Dr. F. J. 
McDonald on ‘“ The Effect of Man on Australian Fauna and Flora”. The 
President thanked the participating speakers and called for a short discussion 
period. 


PROCEEDINGS OF THE LINNEAN SocrETY oF NEw SoutH WALES, VoL. 96, Part 4 


310 ABSTRACT OF PROCEEDINGS 


SPECIAL GENERAL MEETING 
24th November 1971 
Held at the Australian Museum, Sydney, at 7.30 p.m. 
Professor N. G. Stephenson, Vice-President, in the chair. 
Business : 
Professor Stephenson outlined the developments of the past year and gave 
reasons for calling a special General Meeting to discuss the need to widen the 


Society’s Act of Incorporation. He then called on Dr. Vickery, the Honorary 
Treasurer, to explain the proposed changes in detail. 


After discussion, the meeting resolved unanimously to endorse the action 
taken by Council to widen the Society’s powers. 


The meeting closed at 8.25 p.m. 


ORDINARY GENERAL MEETING 
24th November 1971 
Held at the Australian Museum, Sydney, on Wednesday, 24th November 
1971, at 3.30 p.m. 
Professor N. G. Stephenson, Vice-President, in the chair. 


Announcements : 

The Chairman welcomed Mr. H. J. deS. Disney and Dr. C. N. Smithers, 
both new members, to their first General Meeting. 

Myall Lakes Committee. The Chairman drew the attention of members to 
a request for help from the Committee to support the report of the State Planning 
Authority regarding leases in the Myall Lakes area; details were available from 
the Secretary. 

A letter from the “Stop Duffy’s Forest Airport’? Committee asking for 
support in opposing the airport scheme was read to the meeting. 
Papers read (by title only): 

1. Petrology and Stratigraphy of the Brayton District, New South Wales. 
By Re Y. O'Reilly. 
. Check List of Northern Territory Plants. By G. M. Chippendale. 


eI Organic Deposits on Foreshore at North Deewhy. By A. R. H. Martin. 

4. A Brachyopid Labyrinthodont from the Lower Trias of Queensland. 
By A. Howie. 

5. Sedimentation in Jervis Bay. By G. Taylor. 


NOTES AND EXHIBITS 


Mr. D. J. McGillivray spoke on Grevillea rosmarinifolia re-discovered in 
Scotland. 

This species was discovered by Allan Cunningham on the banks of the Cox’s 
River in 1822. Since that time it has not been recorded trom its native habitat, 
nor known from cultivation in Australia. Several closely related species are 
known from New South Wales and Victoria. 

Within a few years of its discovery Grevillea rosmarinifolia was in cultivation 
and flowering in England. It was illustrated in Robert Sweet’s ‘* Flora 
Australasica’’ in 1828, and also in Curtis’s Botanical Magazine in 1872. 

In 1969 aplant of this species was observed by the speaker growing in the open 
beside a building in the Edinburgh Botanic Garden. Although it was not in flower 
and was labelled Grevillea lanata (an error for G. lanigera), it showed a close resem- 
blance to my recollection of the type specimen seen at the Herbarium, Royal Botanic 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH WALES, Vou. 96, Part 4 


ABSTRACT OF PROCEEDINGS oul 


Gardens, Kew. Cuttings were sent from Edinburgh to the Royal Botanic 
Gardens, Sydney, and now a number of thriving plants are in cultivation in 
Sydney. 

Dr. A. Ritchie and Dr. Joan Beattie exhibited a fossil vertebrate collected 
from the rock platform on the northern side of Long Reef immediately to the 
east of the area known as ‘“‘ The Basin” in August this year. The horizon in 
which the specimen occurred was near the top of the Collaroy Claystone, 
Narrabeen Group (Triassic). 

The bones included a femur (about 10 inches long), scapula, rib and other 
fragments and have been provisionally identified as amphibian (labyrinthodont) ; 
their size indicates that the individual was some 12 to 14 feet in length. Dr. 
Anne Howie, who has been working on faunas from the Rewan Formation (of 
equivalent age) of the Bowen Basin, has recently examined this specimen and it 
is hoped that she will, on further examination, find the material sufficient for 
identification. 

Mr. H. J. de S. Disney, Curator of Birds at the Australian Museum, showed 
a map and aerial photograph of Lord Howe Island and also study skins of the 
Flightless Lord Howe Island Woodhen, Tricholimnas sylvestris, with a skin of 
a New Zealand Weka, Gallirallus australis. Also exhibited was the skeleton of 
a Woodhen to show the tiny sternum, not much bigger than that of the now 
extinct Doctor Bird or Vinous-tinted Blackbird, Turdus vinitinctus (also on 
display). 

Mr. Disney said he first visited the island and Mt. Gower in November 1969 
with Dr. Smithers, and found that the Woodhens appeared to be present only 
in very small numbers on top of the mountain. As part of a general survey of 
the flora and fauna of the island a more intense investigation was started in 
February of this year. Birds were caught and colour-banded ; on subsequent 
visits their movements were recorded. From these records, as shown on a map 
of the plateau, there are apparently only 20 birds occupying 10 territories. 


The birds originally were found everywhere on the island. In 1914 they 
were still common in Erskine Valley Little Slope and on Mt. Lidgbird. In 1936 
some were still present in Erskine Valley, but they are now confined to the top 
of Mt. Gower. 

In January 1972 it is hoped that data on breeding success will be obtained ; 
however, there 1s some indication that the summit does not provide an optimum 
habitat. 

The decline in the species may be the result of habitat disturbance by goats 
and pigs, or by predation by rats (which were introduced in 1918) or cats. 

The habits of these birds are similar in many respects to those of the Weka 
of New Zealand. Little is known of their breeding habits, but some have been 
recorded nesting in petrel burrows. 


From the archives of the Australian Museum Mr. G. P. Whitley exhibited 
a photographic portrait of Dr. Haynes Gibbes Alleyne (1815 ?-1882), one of the 
founders of the Linnean Society of New South Wales. 

Dr. Cogger exhibited a living pair of Fijian Iguanas (Brachylophus fasciatus) 
from the island of Kadavu. These lizards are now uncommon in Fiji and are 
regarded as an endangered species. <A closely-allied form occurs in Tonga. 

The family Iguanidae, to which these lizards belong, is otherwise confined 
to the Americas and Madagascar ; its representation in the south-west Pacific 
area is a unique situation in cvartcprane zoogeography, and the Fijian and Tongan 
lizards clearly represent an example of “ rafting” from continental America. 

Dr. C. N. Smithers exhibited specimens of the seven commonest species of 
butterflies from Norfolk Island. Eleven species are known, of which nine occur 


PROCEEDINGS OF THE LINNEAN Society oF NEw SoutH Watgs, Vot. 96, Part 4 


312 ABSTRACT OF PROCEEDINGS 


in Australia. Two of these have recognizable island forms ; the eleventh species 
is known also from New Caledonia. 

A map of Norfolk Island, showing the proposed National Park, was exhibited. 
Action to establish the park is being taken following a report from the Australian 
Conservation Foundation based on a survey carried out in 1967. 

This meeting concluded the session. The Ninety-seventh Annual General 
Meeting, together with the next Ordinary General Meeting, will be held on 
Wednesday, the 29th March 1972. 


PROCEEDINGS OF THE LINNEAN SocreTy or New SourH WaAtss, Vow. 96, Part 4 


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LIST OF MEMBERS 
(31st December, 1971) 


ORDINARY MEMBERS 
(An asterisk (* ) denotes Life Member) 


*Allman, Stuart Leo, B.Sc.Agr., M.Sc., 99 Cumberland Avenue, Collaroy, N.S.W., 2097. 

Anderson, Derek John, Ph.D., Australian National University, P.O. Box 4, Canberra, 
A.C.T., 2601. 

Anderson, Donald Thomas, B.Sc., Ph.D., School of Biological Sciences, Department of 
Zoology, Sydney University, 2006. 

Anderson, Mrs. Jennifer Mercianna Elizabeth, B.Sc.Agr., 24 Watson Street, Bondi, N.S.W.., 


2026. 

Andrew, Mrs. Phillipa Audrey, M.Sc. (née Croucher), 10 Black Street, Watsonia, Victoria, 
3087. 

Ardley, John Henry, B.Sc. (N.Z.), Messrs. William Cooper and Nephews (Australia) Pty. 


Ltd., P.O. Box 12, Concord, N.S.W., 2137. 
*Armstrong, Jack Walter Trench, ‘“ Cullingera’’, Nyngan, N.S.W., 2825. 
Ashton, David Hungerford, B.Sc., Ph.D., 92 Warrigal Road, Surrey Hills, Victoria, 3127. 


Bain, Miss Joan Maud, M.Sc., Ph.D., 10/1 Spencer Road, Killara, N.S.W., 2071. 

Baker, Professor Hldred Percy, B.Sc.Agr., Ph.D., Department of Agricultural Botany, 
Sydney University, 2006. 

Ballantyne, Miss Barbara Jean, B.Sc.Agr., N.S.W. Department of Agriculture, Private 
Mail Bag No. 10, Rydalmere, N.S.W., 2116. 

Bamber, Richard Kenneth, F.S.T.C., 113 Lucinda Avenue South, Wahroonga, N.S.W., 
2076. 

Barlow, Bryan Alwyn, B.Sc., Ph.D., School of Biological Sciences, The Flinders University 
of Scuth Australia, Bedford Park, South Australia, 5042. 

Basden, Ralph, M.Ed., B.Sc. (Lond.), F.R.A.C.I., A.S.T.C., 183 Parkway Avenue, 
Hamilton, N.S.W., 2303. 

Baur, George Norton, B.Sc., B.Sc.For., Dip.For., 3 Mary Street, Beecroft, N.S.W., 2119. 

*Beadle, Professor Noel Charles William, D.Sc., University of New England, Armidale, 


N.S.W., 2350. 
Beard, John Stanley, M.A., B.Sc., D.Phil., Director, Royal Botanic Gardens, Sydney, 
N.S.W., 2000. 


Beattie, Joan Marion, D.Sc. (née Crockford), 2 Grace Avenue, Beecroft, N.S.W., 2119. 

Bedford, Geoffrey Owen, B.Sc., 87 Jacob Street, Bankstown, N.S.W., 2200. 

Bennett, Miss Isobel Ida, Hon.M.Se., School of Biological Sciences, Department of 
Zoology, Sydney University, 2006. 

Bertus, Anthony Lawrence, B.Sc., Biology Branch, N.S.W. Department of Agriculture, 
Private Mail Bag, No. 10, Rydalmere, N.S.W., 2116. 

Besly, Miss Mary Ann Catherine, B.A., School of Biological Sciences, Department of 
Zoology, Sydney University, 2006. 

Bishop, James Arthur, Department of Genetics, The University of Liverpool, Liverpool 3, 
England. 

sinenee John Allan Philip, LL.B. (Syd. Univ.), 25 Holden Street, Ashfield, N.S.W., 
2131. 

Blake, Stanley Thatcher, D.Sc. (Qid.), 1110 Waterworks Road, The Gap, Queensland, 4061. 

Blaxell, Donald Frederick, D.D.A., B.Sc., Royal Botanic Gardens, Sydney, 2000. 

Bourke, Terrence Victor, B.Se.Agr., c/- Department of Agriculture, Stock and Fisheries, 
Popondetta, Papua. 

Boyd, Robert Alexander, B.Sc., Department of Botany, University of New England, 
Armidale, N.S.W., 2350. 

Brett, Robert Gordon Lindsay, B.Sc., 48 Main Road, Lindisfarne, Tasmania, 7015. 

Brewer, Ilma Mary, D.Sc., 13 Wentworth Road, Vaucluse, N.S.W., 2030. 

Briggs, Miss Barbara Gillian, Ph.D., National Herbarium of N.S.W., Royal Botanic 
Gardens, Sydney, 2000. 

Browne, Ida Alison, D.Sc. (née Brown), 363 Edgecliff Road, Edgecliff, N.S.W., 2027. 

Browne, William Rowan, D.Sc., F.A.A., 363 Edgecliff Road, Edgecliff, N.S.W., 2027. 

Burden, John Henry, 1 Havilah Street, Chatswood, N.S.W., 2067. 

*Burges, Professor Norman Alan, M.Sc., Ph.D., Vice-Chancellor, The New University of 
Ulster, Coleraine, County Londonderry, Northern Ireland. 

Burgess, Ian Peter, B.Sc.For., Dip.For., The Forestry Office, Coff’s Harbour, N.S.W., 
2450. 

Burns, James, A.A.S.A., 127 Plateau Road, Avalon Beach, N.S.W., 2107. 

Byrnes, Norman Brice, B.Sc., Dip.Ed., Box 2227, G.P.O., Darwin, Northern Territory, 
5794, Australia. 


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LIST OF MEMBERS 


Cady, Leo Isaac, P.O. Box 88, Kiama, N.S.W., 2533. 

Campbell, Keith George, D.F.C., B.Sc.For., Dip.For., M.Se., 17 Third Avenue, Epping, 
N.S.W., 2121. 

Campbell, Thomas Graham, Division of Entomology, C.S.I.R.O., P.O. Box 109, City, 
Canberra, A.C.T., 2601. 

Canning, Miss Hstelle Margaret, B.Sc. (Melb.), 50 Swan Street, Wangaratta, Victoria, 3677. 

*Carey, Professor Samuel Warren, D.Sc., Geology Department, University of Tasmania, 
Hobart, Tasmania, 7000. 

Carne, Phillip Broughton, B.Agr.Sci. (Melb.), Ph.D. (London), D.1.C., 4 Alt Crescent, 
Ainslie, A.C.T., 2601. t 

Carolin, Professor Roger Charles, B.Sc., A.R.C.S., Ph.D., School of Biological Sciences, 
Department of Botany, Sydney University, 2006. 

Casimir, Max, B.Sc.Agr., Entomological Branch, N.S.W. Department of Agriculture, 
Private Mail Bag, No. 10, Rydalmere, N.S.W., 2116. 

*Chadwick, Clarence Earl, B.Sce., Entomological Branch, N.S.W. Department of Agri- 
culture, Private Mail Bag No. 10, Rydalmere, N.S.W., 2116. 

Chambers, Thomas Carrick, M.Sc. (N.Z.), Ph.D., Botany School, University of Melbourne, 
Parkville, Victoria, 3052. 

Chippendale, George McCartney, B.Sce., 4 Raoul Place, Lyons, A.C.T., 2606. 

Cho, Kai Yip, B.Sc., Ph.D., Department of Microbiology, University of Sydney, N.S.W.., 
2006. 

*Churchward, John Gordon, B.Sc.Agr., Ph.D., “ Erlangga’’, Glen Shian Lane, Mount 
Eliza, Victoria, 3930. 

Clark, Laurance Ross, M.Sc., ¢/- C.S.I.R.O., Division of Entomology, P.O. Box 109, City, 
Canberra, A.C.T., 2601. 


Clarke, Miss Lesley Dorothy, Ph.D., Lot 21, “‘ Bimbadgen”’, Wyoming Road, Dural, 
N.S.W., 2158. 

Clarke, Mrs. Muriel Catherme, M.Sc. (née Morris), 122 Swan Street, Morpeth, N.S.W., 
2321. 


Cleland, Professor Sir John Burton, M.D., Ch.M., C.B.E., 1 Dashwood Road, Beaumont, 
Adelaide, South Australia, 5066. 

Clough, Barry Francis, B.Sc.Agr., School of Biological Sciences, Department of Botany, 
Sydney University, 2006. 

Clyne, Mrs. Densey, 7 Catalpa Crescent, Turramurra, N.S.W., 2074. 

Cogger, Harold George, M.Sc., Ph.D., Australian Museum, P.O. Box A285, Sydney South, 
N.S.W., 2000. 

Colless, Donald Henry, Ph.D. (Univ. of Malaya), c/- Division of Entomology, C.S.I.R.O., 
P.O. Box 109, City, Canberra, A.C.T., 2601. 

Common, Ian Francis Bell, M.A., M.Se.Agr., C.S.I.R.O., Division of Entomology, P.O. 
Box 109, City, Canberra, A.C.T., 2601. 

Conacher, Robert Davy, LL.B., 15 Terrigal Avenue, Turramurra, N.S.W., 2074. 

Cook, Mrs. Margaret Lee (née Debenham), B.Sc., 40/42 Avoca Street, Randwick, N.S.W., 
2031. 

Crawford, Lindsay Dinham, B.Sc., c/- Victorian Plant Research Institute, Department 
of Agriculture, Burnley Gardens, Melbourne, Victoria, 3000. 

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., 2601. 


Dart, Peter John, B.Sc.Agr., Ph.D., Soil Microbiology Department, Rothamsted Expert- 
mental Station, Harpenden, Herts., England. 

Dartnall, Alan John, B.Se., Tasmanian Museum, Box 1164M, G.P.O., Hobart, Tasmania, 
7001. 

Davies, Stephen John James Frank, B.A. (Cantab.), Ph.D., C.S.I.R.O., Private Bag, 
Nedlands, Western Australia, 6009. 

Davis, Professor Gwenda Louise, Ph.D., B.Sc., Faculty of Science, University of New 
England, Armidale, N.S.W., 2350. 


De Nardi, Mrs. Jan Christina, B.Sc. (Q’ld.) (née Morrow), 42 William Street, Paddington, 
N.S.W., 2021. 

Disney, Henry John de Suffren, M.A., The Australian Museum, College Street, Sydney, 
2000. 


Dobrotworsky, Nikolai V., M.Se., Ph.D., Department of Zoology, University of Melbourne, 
Parkville, Victoria, 3052. 

Domrow, Robert, B.A., B.Sc., Queensland Institute of Medical Research, Herston Road, 
Herston, Queensland, 4006. 


Durie, Peter Harold, M.Sc., C.S.I.R.O., Veterinary Parasitology Laboratory, Yeerongpilly, 
Queensland, 4105. 


Dyce, Alan Lindsay, B.Sce.Agr., 48 Queen’s Road, Asquith, N.S.W., 2078. 


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LIST OF MEMBERS oly 


Edwards, Dare William, B.Sc.Agr., Forestry Commission of N.S.W., Division of Wood 
Technology, 96 Harrington Street, Sydney, 2000. 

pees Edward John, B.A., B.Sc., Dip.Ed., 38 Shirlow Avenue, Faulconbridge, N.S.W., 

Endean, Robert, M.Sc., Ph.D., Department of Zoology, University of Queensland, St. 
Lucia, Queensland, 4067. : 

English, Miss Kathleen Mary Isabel, B.Sc., 6/168 Norton Street, Leichhardt, N.S.W., 2040. 


Evans, Miss Gretchen Pamela, M.Sc., 27 Frederick Street, Taringa, Queensland, 4066. 


Facer, Richard Andrew, Department of Geology, Wollongong University College, 
Wollongong, N.S.W., 2500. 

*Fairey, Kenneth David, Box 1176, G.P.O., Sydney, 2001. 

Filewood, Lionel Winston Charles, c/- Department of Agriculture, Stock and Fisheries, 
Konedobu, Papua. 

Flood, Peter Gerard, B.Sc.(Hons.), P.O. Box 52, Spit Junction, N.S.W., 2088. 

Ford, Miss Judith Helen, 57 Boyce Street, Glebe, N.S.W., 2037. 

Fraser, Miss Lilian Ross, D.Sc., 1 Laurence Street, Pennant Hills, N.S.W., 2120. 

Freeman, Brian Warwick, B.Sc., Brain Research Unit, Department of Physiology, Sydney 
University, Sydney, 2006. 


*Garretty, Michael Duhan, D.Sc., Box 763, Melbourne, Victoria, 3001. 
Gerretson-Cornell, Luciano, 1/62 The Kingsway, Cronulla, N.S.W., 2230. 
Goodfellow, David Ralph, 3 Tobruk Avenue, Carlingford, N.S.W., 2118. 


Greenwood, William Frederick Neville, 11 Wentworth Avenue, Waitara, N.S.W., 2077. 


Griffin, David Michael, M.A., Ph.D. (Cantab.), School of Agriculture, Sydney University, 
2006. 
*Griffiths, Mrs. Mabel, B.Sc. (née Crust), 54 Delmar Parade, Dee Why, N.S.W., 2099. 


Griffiths, Mervyn Edward, D.Se., Wildlife Survey Section, C.S.I.R.O., P.O. Box 109, 
City, Canberra, A.C.T., 2601. 

*Gunther, Carl Ernest Mitchelmore, M.B., B.S., D.T.M., D.T.M. & H. (England), M.B.E., 
29 Flaumont Avenue, Lane Cove, N.S.W., 2066. 


Hadlington, Phillip Walter, B.Sc.Agr., 129 Condamine Street, Balgowlah, N.S.W., 2093. 

Hamond, Richard, B.Se., Ph.D., C.S.I.R.O., Box 21, Cronulla, N.S.W., 2230. 

Hannon, Miss Nola Jean, B.Se., Ph.D., 22 Leeder Avenue, Penshurst, N.S.W., 2222. 

Harden, Mrs. Gwenneth Jean, M.Se. (née Hindmarsh), Kellys Plains Road, Armidale, 
N.S.W., 2350. 

Hardwick, Reginald Leslie, B.Sc., Parklands, 12c/1 Evelyn Avenue, Concord, N.S.W., 2137. 

Hayden, Mrs. Elizabeth Jean, B.Sc. (Melb.), 17 Malara Street, Waramanga, A.C.T., 2611. 

Hennelly, John Patten Forde, B.Sc., Highs Road, West Pennant Hills, N.S.W., 2120. 

Hewitt, Bernard Robert, B.A. (Q’ld.), B.Sc. (Syd.), M.Se., (N.S.W.), A.R.A.C.I., Pegawai 
Pertanian, Bandar Seri Begawan, Negri Brunei. 

Hewson, Miss Helen Joan, B.Sc.(Hons.), Ph.D., Department of Botany, School of General 
Studies, Australian National University, P.O. Box 4, Canberra, A.C.T., 2600. 

Higginson, Francis Ross, B.Sc.Agr.(Hons.), Ph.D., Soil Conservation Service of N.S.W., 
Box 4293, G.P.O., Sydney, 2001. 

Hill, Professor Dorothy, Ph.D., D.Sc., F.A.A., F.R.S., Department of Geology and 
Mineralogy, University of Queensland, St. Lucia, Queensland, 4067. 

*Hindmarsh, Miss Mary Maclean, B.Sc., Ph.D., 4 Recreation Avenue, Roseville, N.S.W., 
2069. 

Holland, Ray James Thurstan, M.A. (Syd.), M.A.C.E., ¢/- Sydney Grammar School, 
College Street, Sydney, 2000. 

*Hotchkiss, Professor Arland Tillotson, M.S., Ph.D. (Cornell), Department of Biology, Uni- 
versity 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. 

Hoult, Errol Hubert, B.Sc.(Hons.), Department of Agronomy, University of New England, 
Armidale, N.S.W., 2350. 

Humpbrey, George Frederick, M.Se., Ph.D., C.S.I.R.O. Marine Biological Laboratory, 
Box 21, Cronulla, N.S.W., 2250. 

Hunt, Glenn Stuart, B.Se., Dip.Ed., School of Zoology, University of New South Wales, 
P.O. Box 1, Kensington, N.S.W., 2033. 


Ingram, Cyril Keith, B.A., B.Ee., “La Mancha ”, Mt. Tomah, via Bilpin, N.S.W., 2758. 


Jackes, Mrs. Betsy Rivers, B.Sc., Ph.D. (Univ. Chicago) (née Paterson), 5 Ashton Street, 
Vincent, Queensland, 4814. 


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TG) 7/11 
1930 


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1956 


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1932 
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1971 
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1948 
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LIST OF MEMBERS 


Jacobs, Miss Janice Lorraine, B.Sc., School of Biological Sciences, Department of Botany, 
Sydney University, 2006. 

Jacobs, Surrey Wilfred Laurence, 7 Yarrara Road, Pymble, N.S.W., 2073. 

James, Sidney Herbert, M.Sc., 54 Holmfirth Street, Mt. Lawley, Western Australia, 6050. 

Jamieson, Barrie Gillean Molyneux, B.Sc., Ph.D. (Bristol), Zoology Department, University 
of Queensland, St. Lucia, Queensland, 4067. 

Jancey, Robert Christopher, M.Sc., Ph.D., c/- Department of Botany, University of Western 
Ontario, London, Ontario, Canada. 

Jefferies, Mrs. Lesly Joan, 18 Derby Street, Camberwell, Victoria, 3124. 

Jenkins, Thomas Benjamin Huw, Ph.D., Department of Geology and Geophysics, Sydney 
University, 2006. 

Johnson, Lawrence Alexander Sidney, B.Sc., c/- National Herbarium, Royal Botanic 
Gardens, Sydney, 2000. 

Jolly, Violet Hilary, M.Sc., Ph.D., 2 Hauraki Road, Takapuna, Auckland 9, New Zealand. 

Jones, Edwin Llewelyn, B.A., P.O. Box 196, Leeton, N.S.W., 2705. 

Jones, Mrs. Valerie, National Herbarium, Royal Botanic Gardens, Sydney, N.S.W., 2000. 

Joplin, Miss Germaine Anne, B.A., Ph.D., D.Sc., Department of Geophysics, Australian 
National University, Canberra, A.C.T., 2600. 

Judd, Howard Kenniwell, Minnamurra Falls Forest Reserve, Box 14, P.O., Jamberoo, 
N.S.W., 2533. 


Keast, James Allen, M.Sc., M.A., Ph.D. (Harvard), Professor of Biology, Department of 
Biology, Queen’s University, Kingston, Ontario, Canada. 

Kerr, Harland Benson, B.Sc.Agr., Ph.D., Summer Institute of Linguistics, P.O. Ukarumpa, 
E.H.D., Territory of New Guinea. 

Kesteven, Geoffrey Leighton, D.Sc., c/- Division of Fisheries and Oceanography, 
C.S.I.R.0O., P.O. Box 21, Cronulla, N.S.W., 2230. 

Kindred, Miss Berenice May, B.Sc., Fachbereich Biologie, Universitat Konstanz, 775 
Konstanz, Postfach 733, Germany. 


Lander, Nicholas Sea, B.Sc., National Herbarium, Royal Botanic Gardens, Sydney, 2000. 

Langdon, Raymond Forbes Newton, M.Agr.Sc., Ph.D., Department of Botany, University 
of Queensland, St. Lucia, Queensland, 4067. 

Lanyon, Miss Joyce Winifred, B.Sc., Dip.Ed., 1/20 Clanalpine Street, Hastwood, N.S.W., 
PDR. 

Lawson, Albert Augustus, “‘ Rego House’, 23-25 Foster Street, Sydney, 2000. 

Lee, Mrs. Alma Theodora, M.Sc. (née Melvaine), Manor Road, Hornsby, N.S.W., 2077. 

Lee, Professor David Joseph, B.Sc., School of Public Health and Tropical Medicine, Sydney 
University, 2006. 

Lewis, Mrs. Alison Kay, B.Sc.(Hons.), Dip.Ed. (née Dandie), Flat 1, Albert Road, Strath- 
field, 2135. 

Littlejohn, Murray John, B.Sc., Ph.D. (W.A.), Department of Zoology, University of 
Melbourne, Parkville, Victoria, 3052. 

Lothian, Thomas Robert Noel, Botanic Gardens, Adelaide, South Australia, 5000. 

Lovedee, Miss Lois Jacqueline, B.Sc. (A.N.U.), 11 Dundilla Road, French’s Forest, N.S.W.., 
2086. 

Luig, Norbert Harold, Ph.D., c/- Faculty of Agriculture, Sydney University, 2006. 


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., 2150. 


Macdonald, Colin Lewis, 7 Watford Close, North Epping, N.S.W., 2121. 


MacInnes, Mrs. Christine D. (née Clarke), B.Sc.(Hons.), 71 Macaulay Road, Stanmore, 
2048. 

Macintosh, Professor Neil William George, M.B., B.S., Department of Anatomy, Sydney 
University, 2006. 

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., 2601. 

Maclean, John Leslie, B.Sc.(Hons.), M.Sc., 17 McCawley Street, Watson, A.C.T., 2602. 

*Mair, Herbert Knowles Charles, B.Sc., 111 Deepwater Road, Castle Cove, N.S.W., 2069. 

Marks, Miss Elizabeth Nesta, M.Sc., Ph.D., Department of Entomology, University of 
Queensland, Brisbane, Queensland, 4067. 

Martin, Angus Anderson, B.Sc.(Hons.), Rand, Department of Zoology, University of 
Melbourne, Parkville, Victoria, 3052. 

Martin, Anthony Richard Henry, M.A., Ph.D., School of Biological Sciences, Department 
of Botany, Sydney University, 2006. 


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1955 
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LIST OF MEMBERS 317 


Martin, Helene Alice, Ph.D., School of Biological Sciences, School of Botany, University 
of New South Wales, P.O. Box 1, Kensington, N.S.W., 2033. 

Martin, Mrs. Hilda Ruth Brownell, B.Sc. (née Simons), ¢c/- Mrs. H. F. Simons, 43 Spencer 
Road, Killara, N.S.W., 2071. 

Martin, Peter Marcus, M.Sc.Agr., Dip.Ed., School of Biological Sciences, Department of 
Botany, Sydney University, 2006. 

Mather, Mrs. Patricia (née Kott), M.Sc., Ph.D. (W.A. and Q’ld.), Department of Zoology, 
University of Queensland, St. Lucia, Queensland, 4067. 

McAlpine, David Kendray, M.Sc., Ph.D., Australian Museum, P.O. Box A285, Sydney 
South, N.S.W., 2000. ; 

McCulloch, Robert Nicholson, M.B.E., D.Sc.Agr., B.Se., Cattle Tick Research Station, 
Wollongbar, N.S.W., 2480. 

*McCusker, Miss Alison, M.Sc., 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., 
2089. 

McGarity, John William, M.Sc.Agr., Ph.D., Agronomy Department, School of Rural 
Science, University of New England, Armidale, N.S.W., 2350. 

McGillivray, Donald John, B.Se.For. (Syd.), Dip.For. (Canb.), P.O. Box 107, Castle Hill, 
N.S.W., 2174. 

McKee, Hugh Shaw, B.A., D.Phil. (Oxon.), Service des Eaux et Foréts, B.P. 285, Noumea, 
New Caledonia. 

McKenna, Nigel Reece, Department of Education, Konedobu, Papua. 

Mercer, Professor Frank Verdun, B.Sc., Ph.D. (Camb.), School of Biological Sciences, 
Macquarie University, North Ryde, N.S.W., 2112. 

Messmer, Mrs. Pearl Ray, “Wychwood ”’, 11 Kuring-gai Avenue, Turramurra, N.S.W., 
2074. 

*Meyer, George Rex, B.Sc., Dip.Ed., B.A., M.Ed., Centre for Advancement of Teaching, 
Macquarie University, North Ryde, N.S.W., 2113. 

*Miller, Allen Horace, B.Se., Dip.Hd., School of Teacher Education, Canberra College of 
Advanced Education, P.O. Box 381, Canberra City, A.C.T., 2601. 

Millerd, Miss Alison Adéle, Ph.D., C.S.I.R.O., Division of Plant Industry, P.O. Box 109, 
City, Canberra, A.C.T., 2601. 

Millett, Mervyn Richard Oke, B.A., “ Beeyung’”’, 72 McNicol Road, Tecoma, Victoria, 
3160. 

Milward, Norman Edward, B.Sc.(Hons.), M.Se., Zoology Department, James Cook 
University of North Queensland, P.O. Box 999, Townsville, Queensland, 4810. 

*Moffat, Mrs. Lynette Anne, B.Sc. (née Holder), 48 Rutledge Street, Hastwood, N.S.W., 
2122. 

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., 2601. 

Moore, Raymond Milton, D.Sc.Agr., Division of Tropical Pastures, C.S.I.R.O., Mill Road, 
St. Lucia, Queensland, 4067. 

Morgan, Mrs. Eva, M.Sc., 5317 Borland Road, Los Angeles, California, 90032, U.S.A. 

Moulds, Maxwell Sidney, 14 Chisholm Street, Greenwich, N.S.W., 2065. 

Muirhead, Warren Alexander, B.Sc.Agr., C.S.I.R.O., Irrigation Research Station, Private 
Mail Bag, Griffith, N.S.W., 2680. 

Mungomery, Reginald William, c/- Bureau of Sugar Experiment Stations, 99 Gregory 
Terrace, Brisbane, Queensland, 4000. 

Myerscough, Dr. Peter J., School of Biological Sciences, Botany Building, Sydney 
University, 2006. 


Nashar, Professor Beryl, B.Sc., Ph.D., Dip.Ed. (née Scott), 43 Princeton Avenue, 
Adamstown Heights, N.S.W., 2289. 

Newman, Ivor Vickery, M.Sc., Ph.D., F.R.M.S., F.L.S., 1 Stuart Street, Wahroonga, 
N.S.W., 2076. 

Nicholls, Anthony Oldham, B.Sc., School of Botany, University of Melbourne, Parkville, 


Victoria, 3052. es 
*Noble, Norman Scott, D.Sc.Agr., M.Se., D.I.C., Unit 21, 1 Lauderdale Avenue, Fairlight, 


N.S.W., 2094. 
Noble, Robert Jackson, B.Sc.Agr., Ph.D., 32a Middle Harbour Road, Lindfield, N.S.W., 
2070. 


Nuzum, Miss Caroline Evelyn, Mountain Lagoon Road, Bilpin, N.S.W., 2758. 


O’Farrell, Professor Antony Frederick Louis, A.R.C.Se., B.Sc., F.R.E.S., Department of 
Zoology, University of New England, Armidale, N.S.W., 2350. 
O’Gower, Professor Alan Kenneth, M.Sc., Ph.D., 20 Gaerloch Avenue, Bondi, N.S.W., 2026. 


PROCEEDINGS OF THE LINNEAN SocirETY or NEw SourH Watgs, Vou. 96, Part 4 


1927 
1950 
1952 
1940 
1962 
1957 
1964 


1922 
1962 


1947 


1969 
1969 


1935 
1938 


1967 


1969 


1960 
1962 
1951 
1952 
1953 


1957 
1961 


1968 


1964 
1967 


1946 
1958 
1932 


1967 
1970 


1960 
1962 
1970 


1919 
1965 


1961 
1950 


LIST OF MEMBERS 


Osborn, Professor Theodore George Bentley, D.Sc., F.L.S., 34 Invergowrie Avenue, 
Highgate, South Australia, 5063. 
Oxenford, Reginald Augustus, B.Sc., 10 Greaves Street, Grafton, N.S.W., 2460. 


Packham, Gordon Howard, B.Se., Ph.D., Box 90, P.O., Northbridge, N.S.W., 2063. 

*Pasfield, Gordon. B.Sc.Agr., 8 McDonald Crescent, Strathfield, N.S.W., 2135. 

Payne, William Herbert, A.S.T.C., A.M.I.H.Aust., M.A.P.E., 250 Picnic Point Road, 
Pieniec Point, N.S.W., 2213. 

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., 2601. 

Pedder, Alan Edwin Hardy, M.A. (Cantab.), Ph.D., Institute of Sedimentary and 
Petroleum Geology, 3303 33rd Street, N.W. Calgary 44, Alberta, Canada. 

Perkins, Frederick Athol, B.Sc.Agr., 93 Bellevue Terrace, Clayfield, Queensland, 4011. 

Philip, Graeme Maxwell, M.Sc. (Melb.), Ph.D. (Cantab.), F.G.8., Department of Geology 
and Geophysics, The University of Sydney, Sydney, N.S.W.. 2006. 

Phillips, Miss Marie Elizabeth, M.Sc., Ph.D., Parks and Gardens Section, Department of 
the Interior, Canberra, A.C.T., 2600. 

Phippard, John Harry, B.Pharm., 5/5944 Blaxland Road, Eastwood, N.S.W., 2122. 

Pickard, John, B.Sc.Agr., National Herbarium of N.S.W., Royal Botanic Gardens, Sydney, 
2000. 

Pope, Miss Elizabeth Carington, M.Sc., C.M.Z.S., 8 Loorana Street, East Roseville, N.S.W., 
2069. 

Pryor, Professor Lindsay Dixon, M.Sc., Dip.For., Department of Botany, School of 
General Studies, Australian National University, Box 197, P.O., City, Canberra, 
ANKCHBo, PAROI 

Pulley, Mrs. Jean May, B.Se.Agr., Dip.Ed., 12 Clisby Close, Cook, A.C.T., 2614. 


Quinn, Christopher John, B.Sc.(Hons.), Ph.D., Department of Botany, School of Biological 
Sciences, University of New South Wales, P.O. Box 1, Kensington, N.S.W., 2033. 


Racek, Albrecht Adalbert, Dr.rer.nat. (Brno, Czechoslovakia), School of Biological 
Sciences, Department of Zoology, Sydney University, 2006. 

Rade, Janis, M.Sce., Flat 28a, 601 St. Kilda Road, Melbourne, Victoria, 3000. 

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 1, Kensington, N.S.W., 
2033. 

Ramsay, Mrs. Helen Patricia, M.Sc., Ph.D., School of Biological Sciences, University of 
New South Wales, P.O. Box 1, Kensington, N.S.W., 2033. 

Reye, Eric James, M.B., B.S. (Univ. Q’Id.), Entomology Department, University of 
Queensland, St. Lucia, Queensland, 4067. 

Reynolds, Miss Judith Louise, Ph.D., 118 Homer Street, Earlwood, N.S.W., 2206. 

Richards, Miss Aola Mary, M.Se.(Hons.), Ph.D. (N.Z.), School of Biological Sciences, 
University of New South Wales, P.O. Box 1, Kensington, N.S.W., 2033. - 

Richards, Bryant Neville, B.Se.For., Ph.D. (Q’ld.), 21 College Avenue, Armidale, N.S.W.., 
2350. 

Richardson, Barry John, 12 Bowden Street, Parramatta North, N.S.W., 2150. 

Richardson, Laurence Robert, M.Se., Ph.D., F.R.S.N.Z., 4 Bacon Street, Grafton, N.S.W.. 
2460. 

Riek, Edgar Frederick, B.Se., Division of Entomology, C.S.1.R.0.. P.O. Box 109, City, 
Canberra, A.C.T., 2601. 

Rigby, John Francis, B.Sc., Geological Survey of Queensland, 2 Edward Street, Brisbane, 
Queensland, 4000. 

*Robertson, Professor Rutherford Ness, C.M.G., F.R.S., B.Sc., Ph.D., F.A.A. Master, 

University House, Australian National University, Box 4, Canberra, A.C.T., 2600. 

Rodd, Anthony Norman, Royal Botanic Gardens, Sydney, 2000. 

Ruello, Nick Vincent, B.Se., 1 Terrell Avenue, Wahroonga, N.S.W., 2076. 


Salkilld, Barry William, Dip.Soc.Stud. (Univ. Syd.), 71 Beresford Road, Thornleigh, 
N.S.W., 2120. 

Sands, Miss Valerie Elizabeth, M.Se., Ph.D., P.O. Box 3066, Sherwood, Whangarei, New 
Zealand. 

Satchell, Professor Geoffrey Harold, B.Sc.(Hons.) (Lond.), Ph.D. (Leeds), 7 Bonnefin Road, 
Hunters Hill, N.S.W., 2110. 

*Scammell, George Vance, B.Sc., 7 David Street, Clifton Gardens, N.S.W., 2088. 

Selkirk, David Robert, School of Biological Sciences, Botany Building, Sydney University, 
2006. 

Selwood, Mrs. Lynne, B.Se., Ph.D. (née Bedford), c.o. Mr. R. G. Bedford, 71 Bourke Street, 
Tamworth, N.S.W., 2340. 

*Sharp, Kenneth Raeburn, B.Sc.. Eng. Geology, 8.M.H.l.A.. Cooma, N.S.W., 2630. 


PROCEEDINGS OF THE LINNEAN SociETy oF NEw SoutH WaAtsEs,. VoL. 96, Part 4 


1952 
1949 


1917 
1930 
1940 


1934 


LIST OF MEMBERS 319 


Shaw, Miss Dorothy Edith, M.Sc.Agr., Ph.D., Department of Agriculture, Stock and 
Fisheries, Port Moresby, Papua-New Guinea. 

eae Mrs. Kathleen Margaret, M.Sc., 43 Robertson Road, Centennial Park, Sydney, 

1. : 

Shipp, Erik. Ph.D., 23 Princes Street, Turramurra, N.S.W., 2074. 

Simons, John Ronald, M.Sc., Ph.D., 242 Kissing Point Road, Turramurra, N.S.W., 2074. 

Slack-Smith, Richard J., 7 Kingston Street. Shenton Park, Western Australia 6008. 

Smith, Eugene Thomas, 22 Talmage Street, Sunshine, Victoria, 3020. 

Smithers, Courtenay Neville, Ph.D., Australian Museum, College Street, Sydney, 2000. 

Smith-White, Professor Spencer, D.Sc.Agr., F.A.A., School of Biological Sciences, Botany 
Building, Sydney University, Sydney, N.S.W., 2006. 

South, Stanley A., B.Sc., 47 Miowera Road, Turramurra, N.S.W., 2074. 

Southcott, Ronald Vernon, D.Se., M.D.. D.T.M.& H., F.A.C.M.A., 2 Taylors Road, 
Mitcham, South Australia, 5062. 

Spencer, Mrs. Dora Margaret, M.Sc. (née Cumpston), No. 1, George Street, Tenterfield, 
N.S.W., 2372. 

Staff, Ian Allen, B.Sc., Dip.Ed., Ph.D., Department of Botany, School of Biological Sciences, 
La Trebe University, Bundoora, Victoria, 3083. 

Stanbury, Peter John, Ph.D., The Macleay Museum. School of Biological Sciences, Sydney 
University, 2006. 

Stead, Mrs. Thistle Yolette, B.Sc. (née Harris), 14 Pacific Street, Watson’s Bay, N.S.W., 
2030. 

Stephenson, Neville George, M.Sc. (N.Z.), Ph.D. (Lond.), School of Biological Sciences, 
Department of Zoology, Sydney University, 2006, 

Strahan, Ronald, M.Sc., Director, Taronga Zoo, P.O. Box 20, Mosman, N.S.W., 2088. 

Straughan, Mrs. Isdale Margaret, B.Sc., Ph.D., Allan Hancock Foundation, University of 
Southern California, University Park, Los Angeles, California, 90007, U.S.A. 

Sullivan, George Emmerson, M.Sc. (N.Z.), Ph.D., Department of Histology and 
Embryology, Sydney University, 2006. 

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, 5085. 


Talbot, Frank Hamilton, M.Sc., Ph.D., F.L.S., Australian Museum, P.O. Box A285, Sydney 
South, N.S.W., 2000. 

Taylor, Keith Lind, B.Sc.Agr., c/- C.S.I.R.O., Division of Entomology, Stowell Avenue, 
Hobart, Tasmania, 7000. 

Tchan, Professor Yao-tseng. Dr., es Sciences (Paris), Department of Microbiology, Sydney 
University, 2006. 

Thompson, Mrs. Joy Gardiner, B.Sc.Agr. (née Garden), 10 Alexander Avenue, Mosman 
N.S.W., 2088. 

Thomson, James Miln, D.Sc. (W.A.), Department of Zoology, University of Queensland, 
St. Lucia, Queensland, 4067. 

Thorne, Alan Gordon, B.A., University of New England, Armidale, N.S.W., 2350. 

Thorp, Mrs. Dorothy Aubourne, B.Sc. (Lond.), Ph.D., “Sylvan Close”, Mt. Wilson, 
N.S.W., 2740. 

Thorpe, Ellis William Ray, B.Sc., University of New England, Armidale, N.S.W., 2350. 

Timms, Brian Victor, B.Se.(Hons.), Zoology Department, Monash University, P.O. Box 92, 
Clayton, Victoria, 3168. 

Tindale, Miss Mary Douglas, D.Sc., 60 Spruson Street, Neutral Bay, N.S.W., 2089. 

Tracey, David, B.Sc.(Hons.), Department of Biological Sciences, Stanford University, 
Stanford, California, 94305, U.S.A. 

*Troughton, Ellis Le Geyt, C.M.Z.S., F.R.Z.S., e/- Australian Museum, P.O. Box A285, 
Sydney South, N.S.W., 2000. 

Tucker, Richard, B.V.Sc., Dr.Vet.M., Veterinary School, Department of Veterinary 

Anatomy, University of Queensland, St. Lucia, Queensland, 4067. 


Valder, Peter George, B.Sc.Agr., Ph.D. (Camb.), School of Biological Sciences, Depart- 
ment of Botany, Sydney University, 2006. 

Vallance, Professor Thomas George, B.Sc., Ph.D., Department of Geology and Geophysics, 
Sydney University, 2006. 

Veitch, Robert, B.Sc., F.R.E.S., 24 Sefton Avenue, Clayfield, Queensland, 4011. 

Vickery, Miss Joyce Winifred, M.B.E., D.Sc., Royal Botanic Gardens, Sydney, 2000. 

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., 2033. 

*Voisey, Professor Alan Heywood, D.Sc., 9 Milton Street, Carlingford, N.S.W., 2118. 


PROCEEDINGS OF THE LINNEAN SocrETy oF New SoutrH WaAtgs, Vou. 96, Part 4 


LIST OF MEMBERS 


Walker, Donald, B.Sc., M.A., Ph.D., F.L.S., 18 Cobby Street, Campbell, Canberra, A.C.T., 
2601. 

Walker, John, B.Sc.Agr., Biological Branch, N.S.W. Department of Agriculture, Private 
Mail Bag No. 10, Rydalmere, N.S.W., 2116. 

Walker, John Charles, B.Sc., (Gen. Science), School of Public Health and Tropical 
Medicine, Sydney University, N.S.W., 2006. 

Walkom, Arthur Bache, D.Sc., 5/521 Pacific Highway, Killara, N.S.W., 2071. 

Wallace, Murray McCadam Hay, B.S8c., C.S.I.R.O., W.A. Regional Laboratory, Nedlands, 
Western Australia, 6009. 

Ward, Mrs. Judith, B.Sc., 16 Mortimer Avenue, New Town, Hobart, Tasmania, 7008. 

Warren, Dr. Alison Anne, B.Sc., Ph.D. (Cantab.) (née Howie), Department of Geology, 
University of Melbourne, Parkville, Victoria, 3052. 

Wass, Robin Edgar, B.Se.(Hons.) (Q’ld.), Ph.D., Department of Geology and Geophysics, 
Sydney University, 2006. 

Waterhouse, Douglas Frew, C.M.G., D.Sc., F.R.S., C.S.I.R.O., P.O. Box 109, City, 
Canberra, A.C.T., 2601. 

*Waterhouse, John Teast, B.Sc., School of Biological Sciences, University of New South 
Wales, P.O. Box 1, Kensington, N.S.W., 2033. 

Watson, Professor Irvine Armstrong, Ph.D., B.Sc.Agr., Faculty of Agriculture, Sydney 
University, 2006. 

Webb, Mrs. Marie Valma, B.Sc., 30 Talfourd Street, Glebe, .N.S.W., 2037. 

Webby, Barry Deane, Ph.D., M.Sc., Department of Geology and Geophysics, Sydney 
University, 2006. 

Webster, Miss Elsie May. 

Wharton, Ronald Harry, M.Sc., Ph.D., Division of Entomology, C.S.I.R.O., Long Pocket 
Laboratories, Indooroopilly, Queensland, 40€8. 

*Whitley, Gilbert Percy, F.R.Z.S., Australian Museum, P.O. Box A285, Sydney South, 
N.S.W., 2000. 

Whitten, Maxwell John, Ph.D., C.S.I.R.O., Division of Entomology, P.O. Box 109, City, 
Canberra, A.C.T., 2601. 

Wildon, David Conrad, B.Sc.Agr., Box 108, Rozelle, N.S.W., 2039. 

Williams, John Beaumont, B.Sc., University of New England, Armidale, N.S.W., 2350. 

Williams, Mrs. Mary Beth, B.Sc. (née Macdonald), 902D Rockvale Road, Armidale, N.S.W., 
2350. 

Williams, Neville John, B.Sc., 56 Karimbla Road, Miranda, N.S.W., 2228. 

Williams, Owen Benson, M.Agr.Sc. (Melbourne), c/- C.S.I.R.O., The Ian Clunies Ross 
Animal Research Laboratory, P.O. Box 144, Parramatta, N.S.W., 2150. 

Willis, Jack Lehane, M.Se., A.A.C.I., 26 Inverallan Avenue, Pymble, N.S.W., 2073. 

Winkworth, Robert Ernest, Rangelands Research Unit, C.S.I.R.O., P.O. Box 109, Canberra 
City, A.C.T., 2601. 

Wood, Alec Edward, B.Sc.Agr., Ph.D., 25 George Street, Bexley, N.S.W., 2207. 

Woodward, Thomas Hmmanuel, M.Sc. (N.Z.), Ph.D. (Lond.), D.I.C., Department of 
Entomology, University of Queensland, St. Lucia, Queensland, 4067. 

Wright, Anthony James Taperell, B.Sc., Department of Geology, Wollongong University 
College, Wollongong, N.S.W., 2500. 


Yaldwyn, John Cameron, Ph.D. (N.Z.), M.Sc., Dominion Museum, Private Bag, Wellington, 


New Zealand. 
Young, Graham Rhys, 8 Spark Street, Earlwood, N.S.W., 2206. 


CoRRESPONDING MEMBERS 
Burgess, Rev. Colin, 2 Hamilton Row, Yarralumla, A.C.T., 2600. 
Jensen, Hans Laurits, D.Sc.Agr. (Copenhagen), State Laboratory of Plant Culture, 
Department of Bacteriology, Lyngby, Denmark. 
Moore, Kenneth Milton, Cutrock Road, Lisarow, N.S.W., 2251. 


PROCEEDINGS OF THE LINNEAN Society or New SoutH WALES, Vou. 96, Part 4 


LIST OF PLATES 321 


PROCEEDINGS 1971 


I.—Lord Howe Island. 
IJ.—Lizards from Lord Howe Island. 
III.—Species of Patiriella. 
IV.—Type localities of Patiriella. 
V-VI.—Late Devonian conodonts. 
VII.—Bacterial leaf spot of zinnia. 
VIII.—Species of zinnia negatively stained. 
IX.—Holotype of Lophopetalum arnhemicum N. Byrnes. 
X-XI.—Ustilago cyperi—lucidi. 
XII-XITI.—Ramularia pitereka on Hucalyptus spp. 
XIV-XV.—Brachyops allos: views of skull. 
XVI.—Rock formations in the Brayton District, N.S.W. 


LIST OF NEW SPECIES 


VoL. 96 

New Species Page 
allos (Brachyops) .. oe 0) obscura (Patiriella) .. Seay 
alpina (Allwaudomyia) : a AG personata (Alluaudomyia) .. 133 
appendiculata (Alluaudomyia) .. 151 pitereka (Ramularia) v7. 08 
arnhemicum (Lophopetalum) oo HD) pseudoexigua (Patiriella) .. 43 
bicornis (Alluaudomyia) .. oo D9 reyel (Alluaudomyia) anal a 
fragmentum (Alluaudomyia) ee eek63 unguistyla (Alluaudomyia).. 139 
fumosipennis (Alluwaudomyia) eles) varia (Allwaudomyia) ge a 
kangarooensis (Nunciella) . . eS verecunda (Alluaudomyia) 143 


New Subspecies 


annulipes australiensis (Alluau- 
domyia) .. a ai alta 


New Combinations 


badia (Nuneciella) .. a eo OO involucrata (Borreria) egiZ0g 
dentata (Nwunciella) ae a: 58 schultzii (Canthium) .. 209 


Index 


Page 

Abstract of Proceedings oo LBS) 

g. Alluaudomyia Kieffer in Australia and 
New Guinea ae 128 

Annual General Meeting. . 6 bi 1 

Aphids of Papua and New Guinea: An 
annotated check list .. 186 

Australian Ceratopogonidae Gointera! 
Nematocera). Part XV: The genus 
Alluaudomyia WKieffer in Australia 
and New Guinea 128 

Authors, notes and instructions 331 

Australian sea stars of the genus Patiriella 
(Asteroidea, Asterinidae) 39 

Bacterial Leaf Spot of Zinnia in New 
South Wales .. : 81 

Balance Sheets oe a 5 

Bertus, A. L., see Walker, J., and 
Bertus, A. L. 

Bertus, A. L., and Hayward, A. C., 
Bacterial Leaf Spot of Zimnia in 
New South Wales 81 

Brachyopid labyrinthodont nae ig 
lower Trias of Queensland . 268 

Brayton District, New South Wales, 
petrology and stratigraphy 282 

Broadbent, P., Fraser, L. R., and Water- 
worth, Y., The reaction of seedlings 
of Citrus spp. and related genera to 
Phytophthora citrophthora 119 

Byrnes, N., A new Species of Beane: 
petalum Caen: a Genus new 
for Australia . 85 

Carolin, R. C., The Trichomes of the 
Goodeniaceae : 8 

Chippendale, G. M., Cheek List of 
Northern Territory Plants 207 

Cogger, H. G., The Ron of Lord 
Howe Island 23 

g. Ctenodiscus, Sea Star Hee Tasmania... 50 

Dartnall, A. J., Australian Sea Stars 
of the genus Patiriella (Asteroidea, 
Asterinidae) 39 

Debenham, Margaret L., Australasian 
Ceratopogonidae (Diptera, Nema- 
tocera). Part XV: The genus 
Alluaudomyia Kieffer in Australia 
and New Guinea a .. 128 

Depositional environment of the organic 
deposits on the foreshore at North 
Deewhy, New South Wales 278 

Elections 4 

Fraser, L. R., see Broadbent, P., Fraser, 


L. R., and Waterworth, Y. 


1971 


Page 

Goodeniaceae, the trichomes of. . . 8 

Grasslands, phenology of species common 
to three semi-arid : 193 

Hayward, A. C., see Bertus, A. L., bes 
Hayward, A. C. 

Howie, Anne, A_ brachyopid laby- 
rinthodont from the lower Trias of 
Queensland BAe . 268 

Hunt, G. 8., The genus Ieporeitio meee 
(Opiliones, Laniatores) with descrip- 
tion of a new species from eee 
Island, South Australia 53 

Jackson, J. H., see Philip, G. M., and 
Jackson, J. H. 

Jacobs, §$., Systematic position of the 
genera J'riodia R.Br. and Plec- 
trachne Henr. (Graminaceae) 175 

Jervis Bay, sedimentation in . 297 

Kangaroo Island, South Australia, g. 
Nunciella Roewer any 
Laniatores) Be 53 

Lamb, K. P., Aphids of Papua eee New 
Guinea: An annotated check list . 186 

Late Devonian Conodonts from ste 
Luton Formation, Northern New 
South Wales .. : 66 

Linnean Macleay Fellowship 3 

Linnean Macleay Lectureship in Micro- 
biology 7H + 

Lophopetalum (Celastraceae), a new 
species of a genus new for Australia 85 

Lord Howe Island, the reptiles of 23 

Maclean, J. L., The Food and Feeding of 
Winter Whiting (Silago maculata 
Quoy and Gaimard) in Moreton Bay 87 

Martin, A. R. H., The depositional 
environment of the organic deposits 
on the foreshore at North go tia 
New South Wales 278 

Members, List of aa 5 > als} 

New Guinea Highland Dee (Canis 
hallstromi), the early history and 
relationships aes $0 oo) (OB 

Northern Territory Plants, check list .. 207 

Notes and Exhibits: 

Cogger, H. G.—Fijian Iguanas .. 311 
Disney, H. J. de 8.—Flightless Lord 
Howe Island Woodhen . 311 
McGillivray, D. J.—Grevillea ros- 
marinifolia re-discovered in 
Scotland : ar .. 310 
Ritchie, A., and Beattie, Joan— 
Fossil vertebrate from Long 
Reef, Collaroy Claystone do ail 


INDEX 


Notes and Exhibits—continued 


Page 
Smithers, C. N.—Butterflies from 
Norfolk Island S43 oc lil 
Whitley, G. P.—Dr. ee Gibbs 
Alleyne .. . 311 
Notes and instructions for authors .. 331 
g. Nunciella Roewer (Opiliones, Lania- 
tores) with description of a new 
species from Kangaroo Island, 
South Australia me ye OS 
O’Reilly, Suzanne Y., Petrology and 
stratigraphy of the Brayton District, 
New South Wales .. 282 
g. Patiriella (Asteroidea, aes) 
Australian sea stars.. 66 do BY) 


Petrology and stratigraphy of the 
Brayton District, New South Wales 282 


Plates, List of 


Philip, G. M., and Jackson, J. H., Late 
Devonian Conodonts Prom the Luton 
Formation, northern New South 
Wales .. Be . 66 


Phytophthora Fie ie: reaction 


of seedlings of Citrus SEP: and 
related genera to so de) 
Reptiles of Lord Howe Island .. 50 2B 


Report on the Affairs of the Society 1 
Science Centre .. os ee 50 Ah 8 


Sea Star of Genus Ctenodiscus from 
Tasmania — oe op so OW 


323 


Page 


Sedimentation in Jervis Bay f 
Shoot Blight of Hucalyptus spp. caused 
by an _ undescribed Ponge of 
Ramularia 
Tasmania, a Sea Star of genus Cionadiseis 
Taylor, G., Sedimentation 
Bay . 5 oe 2% : 
Triodia R.Br. and Plectrachne Henr. 
(Gramineae), systematic position of 


Troughton, Ellis, The early history and 
relationships of the New Guinea 
Highland Dog (Canis hallstrom1) 


Ustilago on Cyperus lucidus R.Br. in 
Australia, an undescribed species of 


Walker, J., An undescribed species of 
Ustilago on Cyperus lucidus R.Br. 
in Australia. With Robert Brown’s 
type collection of C. lucidus. . 


Walker, J., and Bertus, A. L., Shoot 
Blight of Hucalyptus spp. caused by 
an undescribed species of Ramularia 


Waterworth, Y., see Broadbent, P., 
Fraser, L. R., and Waterworth, Y. 


Williams, O. B., Phenology of species 
common to three semi-arid ies 
lands 


in Jervis 


Winter Whiting (Szllago Gan ques 
and Gaimard) in Moreton Bay, food 
and feeding oe : 


Zinnia in New South Wales, a Bacterial 
Leaf Spot .. ye ee a 


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