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THE

RMRKOCE EW NGS

OF TILK

LISMEAN SOCIETY @

i el

OF

Nise Sours WWONES

FOR THE YEAR

WO 0

VOL. LIT.

WITH FIFTY PLATES
and 372 Text-figures.

SYDNEY:
PRINTED AND PUBLISHED FOR THE SOCIETY BY
THE AUSTRALASIAN MEDICAL PUBLISHING CO., LTD.,
Seamer Street, Glebe, Sydney,
and
SOLD BY THE SOCIETY.
1927-1928.

CONTENTS OF PROCEEDINGS, 1927.

PART I (No. 210).
(Issued 15th April, 1927.)

Pages

Presidential Address delivered at the Fifty-second Annual Meeting,
30th March, 1927, by E. W. Ferguson, M.B., Ch.M., D.P.H. a raeturteyts i-xxviii
Elections and Announcements .. .. PB Poe! Wren bne XXViii
Balance Sheets for the Year ended 31st Tacomber: 1926 SA Cle Ges mer 1 ee0.9D.05-5:0.0]
Joseph James Fletcher RMT SEL ENN Maia eke pi Mis Ae iee bars inate git cera use h ery AM ORR NT LL CLD

PART II (No. 211).
(Issued 11th July, 1927.)

Notes on Australian Diptera. No. x. By J. R. Malloch. (Communicated
by Dr. HE. W. Ferguson.) (Twelve Text-figures.) .. .. .. .. .. 1- 16
The Influence of certain Colloids upon Fermentation. Parts iv-vi. By
R. Greig-Smith, D.Sc., Macleay Bacteriologist to the Society. (Two
Text-figures. ) ER ee Lesage 7 Pender A ta) comets 17- 24
A Note on Reproductive Pitansiene in some Lizard? By H. Claire
Weekes, B.Sc., Linnean Macleay Fellow of the Society in Zoology. -°
(Three Text-figures. ) Eb gates Ce, ent ee ae agent em are 25- 32
Notes on Australian Mosquitoes (Dinterat Culicidae). Part i. The
Anophelini of the Mainland. By I. M. Mackerras, M.B., Ch.M., B.Sc.,
Linnean Macleay Fellow of the Society in Zoology. (Three Text-
figures. ) cia hEAaS ; 33- 41
The Interpretation of the Radial Field be tne Wing in ne IN Gamibectous
Diptera, with Special Reference to the Tipulidae. By Charles P.
Alexander, Ph.D. (Communicated by Dr. EH. W. Ferguson). (Ninety-

two Text-figures.) Face eine Se ree y Nene cect aoe fovea Dade seus 42- 72
The Anatomy of Cheilanthes Delee: By May M. Williams, M.Sc., Linnean

Macleay Fellow of the Society in Botany. (Twenty Text-figures.) .. 73-. 84
The Geology of the Country between Lamb’s Valley and the Paterson

River. By G. D. Osborne, B.Sc. (Plate i and two Text-figures.) .. 85-103
A New Deltopecten from the Illawarra District, N.S.W. By John Mitchell. j

(CRIA TC RITE) etamete eer in crmeml crept senmenreC oe ct are Vater cote be a eal Ree 104
The Fossil Estheriae of Australia. Pantene: By John Mitchell.

(Plates ii-iv.) SP ahaa re ldcaage 105-112
A Note on a Dicotyledonous Fossil ood fom Ulladulla, New South

Wales. By C. Barnard, B.Se. (Plates v-vi and six Text-figures.) .. 113-121

On a Case of Natural Hybridism in the Genus Grevillea (N.O. Proteaceae).
By C. T. Musson and the late J. J. Fletcher, M.A., B.Sc. (Plate vii.) 122-132
The Physiographic and Climatic Factors controlling the Flooding of the
Hawkesbury River at Windsor. By Lesley D. Hall, B.Se. (Plate viii
and nine Text-figures.) SC TARSTAT MERAY Recta eRe Ae Noy VEN Be ey Boe Lao be

309812

iv. CONTENTS.

New Gall-forming Thysanoptera of Australia. By Dudley Moulton.
(Communicated by W. W. Froggatt.) (Plate ix and fourteen Text-
figures. ) . Biker Pncamd Os cee coe Stee

An Ecological Study of ine igen GE Mount Wilson. Part iv. Habitat
Factors and Plant Response. By John McLuckie, M.A., D.Sc., and
Arthur H. K. Petrie, M.Sc.. (Twenty Text-figures.)

Two New Species of Setaria from Western Australia. By Dr. aA S,
Hitchcock. (Communicated by W. M. Carne.)

SEVACEU Ty olelstin NO 2i'2))):
(Issued 25th October, 1927.)

The Vegetation of the Kosciusko Plateau. Part i. The Plant
Communities. By J. McLuckie, M.A., D.Sc., and A. H. K. Petrie, M.Sc.
(Plates x-xix and eight Text-figures.) SE AE Aad Siac Seah

Australian Coleoptera: Notes and New Species. No. v. By H. J. Carter,
B.A., F.E.S. (Six Text-figures.) Pca Tears iho ES eet WN AG

The Gasteromycetes of Australasia. vii. The Genera Disciseda and
Abstoma. By G. H. Cunningham. (Plates xx-xxii.)

The Gasteromycetes of Australasia. viii. The Genus Mycenastrum. By
G. H. Cunningham. (Plate xxii, figs. 5-8.)

The Gasteromycetes of Australasia. ix. Keys to the Genera andl Snecies
of the Lycoperdaceae. By G. H. Cunningham .. sige wae

The Xerophytic Structure of the Leaf in the Australian Pecteccene: Part i.
By A. G. Hamilton. (Plates xxiii-xxv and twenty-seven Text-figures. )

Australian Hesperiidae. Part i. Notes and Descriptions of New Forms.
By G. A. Waterhouse, D.Sc., B.E., F.E.S. (Plate xxvi.)

Notes on Australian Mosquitoes (Diptera-Culicidae). Part ii. The
Zoogeography of the Subgenus Ochlerotatus, with Notes on the
Species. By I. M. Mackerras, M.B., Ch.M., B.Sc., Linnean Macleay
Fellow of the Society in Zoology. (Hight Text-figures.) eh ations

Notes on Australian Diptera. No. xi. By J. R. Malloch. (Communicated
by Dr. I. M. Mackerras.) (Twenty-three Text-figures. ) :

Notes on Australian Diptera. No. xii. By J. R. Malloch. (Communicated
by Dr. I. M. Mackerras.) (Two Text-figures.) .. BRU! Vie ee lier

Descriptions of New Species of Australian Coleoptera. Part xix. By
A. M. Lea, F.E.S. sveas estan Totals «ster Seasys

Additional Flora of fhe Gambari Plateau, 1926. "By HK. C. Chisholm,
M.B., Ch.M.

On the Phylogeny of some “Diptere: SBrachvecea: Ba G. H. Hardy. (Two :

Text-figures. ) 7 :
Further Notes on a New Giaeuincation of AYetealiad Robbersies (piptere:
Asilidae). By G. H. Hardy. (One Text-figure.)

PART IV (No. 213).
(Issued 15th December, 1927.)

Notes on Australian Diptera. No. xiii. By J. R. Malloch. (Communicated
by Dr. I. M. Mackerras.) (¥ifteen Text-figures.)

Pages.

153-160

161-184

185-186

187-221

222-234

235-244

245-246

247-257

258-274

275-283

284-298

299-335

336-353

354-377

378-379

380-386

387-398

399-446

CONTENTS. We

Pages.
Notes on Australian and Exotic Sarcophagid Flies. By G. H. Hardy.

(Eleven Text-figures. ) Sh!) TEE ee Be) Diag wea ey BNC. oP aa -459
Notes on Australian Marine Algae. iv. The Australian Species of the
Genus Spongoclonium. By A. H. S. Lucas, M.A., B.Se. (Plates

SO QRIED OO Cs no INS cabe IRAE MIG Olay cro’ ava eth ow to AS Os, aumee tO ec TAU
Studies in the Goodeniaceae. i. The Life-history of Dampiera stricta
(R.Br.). By P. Brough, M.A., B.Sc., B.Se.(Agr.). (Plates xxxvi-xxxvii

and fifty-eight Text-figures.) .. .. Svat | eR aduekey ey RO aL ERT Gh Oe) we EEL 498
Placentation and other Phenomena in ihe Scincid Lizard, Lygosoma
. (Hinulia) quoyi. By H. Claire Weekes, B.Sc., Linnean Macleay Fellow
of the Society in Zoology. (Plates xxxviii-xl and twenty-three Text-

figures. ) Bik Oe OLR Ee: ON LIE ite MOE ee
Notes on Australian Marine Aleaes v. By A. H. S. Lucas, M.A., B.Sc.
@Blatesw xlizxivilig ss. 555-562

Mosquito Control in the Municipality, of cane Cove: New South Wales.
By B. Bertram. (Communicated by Dr. I. M. Mackerras.) (Plates

xlix-l, and one Text-figure.) aig ne See te ao 0o-D09
A New Dendrobium for New South Wales ead DiceeIond By Rev.
H. M. R. Rupp. (Four Text- -figures. ) Sais Ree iss 0 tats aychod ee inet al rake. SD OSDIEL

PART V (No. 214).
(Issued loth February, 1928.)

ADStract OM PhOCCCHIME Sica mein fiat Mean him cat Nie loose Ak apres SU ha xliii-liv
Donations#andsHxchansesm enna 5) Winey mt en tenn eT On aunt a tapes lv-lxxiii
List of Members Ooo NEG) erro ne WenGDL Monee Gipsy motets tt Gar pts eb oO-chelb.o-ayvbi
Index

1xxix-xcVv

vi. CONTENTS.

LIST OF NEW TRIBAL AND GENERIC NAMES PROPOSED IN THIS
VOLUME (1927).

Page.
Anthracomyia (Callophorinae, Diptera) .. .. .. .. .. .. .. .. pas Galt)
Apinocis (Curculionidae) Maen Ng Ute Sn ae Oise Seraph RSet pay Vena yc Ramm PLE
Asilini (Asilinae) Rb Bae tae ie ake eikrc rete -aae Uicaetinrs a i DR cacao ta Sian Nd else BIEN)
Choleothrips (Kladothripinae) Bye eC eM ari tae A Dr Re etc aa AU lel ore ae Ngan! TILER
IHOKCOM SON as (COMBO oboe, IDEN) 56 oo Go Go lod oo of bo oo 7H
IHUSRCOTON IG (CMOS ioMmuoT, IDE) og bo of 05 oo vou. oo ‘o0 co, oo Gai
EGDRTANUSOASIINAC)) ais RR pe eR, ere ree te use aca: oc eG Wines em OO
ILEMIGOOaPIS (CURGWTOIICI®)) oo “oo \ido. Ge oe oo do OB oo 00, 66 “oo B70
Melanina (Sapromyzidae) Seems, ease, Weegee | Ries ect Mii Ses Vr at ele RET
Ommatiini (Asilinae) OAT Uehara itis Ma inte COs OR et ee Me. SRD OS RU a i ee SOLO)
LUTE OGS. (AEXOUE TO ONIGENE Arcee” ae a ou hub) ERA Dome vcig NEpou Memos COG
SEMtUSU LUO CDIDECK AN meas am cee puta any lees Uaten costeeeanye CP aiNCRN I Out ar mann)

LIST OF PLATES.
PROCEEDINGS, 1927.

i.—Geological Map of Lamb’s Valley-Paterson District.
ii.—Deltopecten rienitsi and species of Estheria.
lii-iv.—Species of Hstheria.

v-vi.—Fossil wood from Ulladulla, N.S.W.

vii—Species of Grevillea.

viii.—Cracks in flood silt deposits.

ix.—Kladothrips, Choleothrips and Dolerothrips.
x.—Topographic map of the Kosciusko Plateau.
xi-xix.—Vegetation, etc., at Mt. Kosciusko.
xXx-xxii.—Australasian species of Disciseda, Abstoma and Mycenastrum.
xxili-xxv.—Sections of leaves of Hakea.
xxvi.—Australian Hesperiidae.

XXVil-xxxiv.—Species of Spongoclonium.

xxxv.—Species of Callithamnion.
XXXV1-XXxVili—Dampiera stricta.
XXXVili-xl.—Placentation in Lygosoma (Hinulia) quoyi.
xli—Codium difforme (?), C. globosum and Avrainvillea erecta.
xlii-xliii—Species of Caulerpa.

xliv.—Hormosira articulata.

xly-xlviii—Species of Chondria.

xlix-l—Views in the Municipality of Lane Cove.

CORRIGENDA.
Page 194, line 19, for sanguisorba, read sanguwisorbae.
Page 562, line 15, for coerulescens, read caerulescens.

£4 Gi Oy gh : ‘ ral

THE

PROCEEDINGS

OF THE

TSN EAN SOCin RY

OF

News Sowrd, | VW ALES

.

WEDNESDAY, 30TH MarcH, 1927.

The Fifty-second Annual General Meeting, together with the Ordinary Monthly
Meeting, was held at Macleay House, 16 College Street, Sydney, on Wednesday
evening, 30th March, 1927.

ANNUAL GENERAL MEETING.

Mr. A. G. Hamilton, Vice-President, in the Chair.
The Minutes of the preceding Annual General Meeting (31st March, 1926)
were read and confirmed.

A letter of apology from the President (Dr. E. W. Ferguson) for his unavoid-
able absence through illness was read.

A letter of greeting to the meeting, dated Cambridge, England, 22nd February,
1927, was read from the Secretary, Dr. A. B. Walkom.

PRESIDENTIAL ADDRESS.

The past year has been one of more than usual activity in scientific affairs
in Australia: The Australasian Association for the Advancement of Science
and Australian National Research Council held their meetings in Perth in August
last; quite a number of Australians have attended the third Pan-Pacific Science
Congress in Tokio, and that important body “the Council for Scientific and
Industrial Research’? has been constituted. The activity of members of the
Society in conducting their own lines of research is emphasized by the fact that
the volume of proceedings was larger than usual and, in addition, there were
eleven papers in hand for 1927 before the close of 1926.

The Australasian Association for the Advancement of Science held its 15th
meeting in Perth during August, 1926. This was the first occasion on which the
Association had ventured to hold its meeting at Perth, some doubt having been
felt as to the possibility of holding a successful meeting in a centre which has

been considered so remote. The unparalleled success of the 18th meeting in every
A

ii. PRESIDENTIAL ADDRESS.

respect, shows that such doubt need no longer prevent Perth taking its place
among the regular centres at which meetings are held.

It was with very great regret that we learnt of the death a few months ago
of the President of the Association, Edward Henry Rennie, M.A., D.Sec., Professor
of Chemistry in the University of Adelaide, who was a native of Sydney, a
graduate of the University of Sydney, where he obtained his M.A. in the same
year (1876) as the late J. J. Fletcher. Professor Rennie also received the B.Sc.,
from the University of London in the same year (1879) as the late J. J. Fletcher.

The Third Pan-Pacific Congress was held in Tokio, Japan, in November, 1926,
under the auspices of the National Research Council of Japan and was attended
by upward of 130 delegates from overseas. The countries represented were those
bordering or having interest in the Pacific and included the United States of
America, Australia, Canada, France, Great Britain, Hawaii, Japan, Netherlands,
Netherlands East Indies, New Zealand, Philippine Islands, and for the first time,
China and Russia.

The subjects discussed at the Congress included all branches of physical and
biological Sciences and a very full programme was carried out.

At this meeting a Pacific Science Association was formed and a constitution
adopted under which future Pacific Science Congresses will be held at intervals
of not less than two years nor more than five years. The next meeting is to be
held in Java in 1929.

Amongst the delegates who attended were the following members of this
Society: Mr. R. H. Cambage, C.B.E., President of the Australian National Research
Council, Messrs. E. C. Andrews and G. H. Halligan and Professors L. A. Cotton,
T. Griffith Taylor and W. N. Benson (New Zealand).

The Great Barrier Reef Committee have completed their boring operations at
Michaelmas Reef, Oyster Cay, near Cairns. The boring party landed during
April, 1926, and operations ceased about the end of August. The boring results
have yielded a rich harvest of information and have shown that the total thickness
of the coral at Oyster Cay is about 427 feet. The material on which the coral
rests is quartz sand, coloured green by the presence of glauconite and containing
fragments of shells of molluscs and of foraminifera. The nature of the underlying
quartz sand suggests that it was formed in shallow water conditions. This quartz
sand was penetrated to a depth of 600 feet and practically all coralline material
above it was loosely coherent, necessitating the use of casing right through. During
the year several pieces of physiographical and geographical work had been
completed and a report on the Barrier Reef Plankton Collections had been prepared.
The Committee has decided to assist the expedition which will be sent cut during
the coming year by the University of Cambridge under the leadership of Mr. fF. A.
Potts, M.A. This expedition proposes to study the marine zoology and botany
of a portion of the Reef.

A highly successful appeal for funds to institute Cancer Research at the
University of Sydney has brought the large sum of over £127,000.

As members already know, the Council decided to perpetuate the memory
of the late J. J. Fletcher by the institution of an annual “J. J. Fletcher Memorial
Lecture”. The lecture, which is to be limited to some branch of natural history,
is to be delivered at the invitation of the Council, and for the first lecture, in
1928, I am sure all will agree that no more fitting choice could have been made
than Sir Baldwin Spencer, one of Mr. Fletcher’s oldest and closest friends in
Australia, who has kindly consented to deliver the lecture early in 1928. In

PRESIDENTIAL ADDRESS. ili.

addition, it is intended to erect a small tablet in memory of Mr. Fletcher as soon
as the opportunity offers.

The concluding part of Volume LI of the Society’s proceedings has been
issued. The complete volume (657 plus xciv pages, 48 plates and 227 text-figures)
contains thirty-five papers from twenty-nine authors, including several representing
first contributions by some of the younger members of the Society. The Society’s
research staff contributed four papers.

Exchange relations with scientific societies and institutions were normal,
the receipt for the session numbering 1,821, as compared with 1,409, 1,457 and 1,450
for previous sessions. The year has been marked by the large number of applica-
tions received from Societies and Institutions in other parts of the world to be
placed on our exchange list. As many as possible of these have been acceded to,
and the following names have been added to the exchange list during the year:
Biologische Wolga-Station, Saratow, Russia; Botanic Gardens, Rio de Janeiro;
Botanical Laboratory of the University of Montreal; College of Agriculture, Los
Banos, Philippine Islands; Dominion Museum, Weliington, New Zealand; Institut
des récherches biologiques a l'Université de Perm, Russia; Marine Biological
Laboratory, Woods Hole, U.S.A.; Siberian Institute of Agriculture and Forestry,
Omsk; and Instituto Botanico, Universidade de Coimbra, Portugal.

The vacancies on the Council caused by the resignations of Mr. J. H.
Campbell and Professor H. G. Chapman, and the death of Mr. J. J. Fletcher were
filled by the election of Mr. A. H. S. Lucas, Professor A. N. Burkitt and Dr. C.
Anderson respectively.

As a result of representations made by Professor Sir Edgeworth David and
Professor A. C. Seward of The University of Cambridge, the International Educa-
tion Board (founded by John D. Rockefeller, Jr. in 1923) awarded to Dr. A. B.
Walkom a Fellowship in Science to enable him to pursue his studies in palaeo-
botany under Professor Seward during 1927. The Council has granted Dr. Waikom
twelve months’ leave of absence to allow him to take advantage of this award;
and has appointed Dr. G. A. Waterhouse as Acting-Secretary during Dr. Walkom’s
absence.

I have much pleasure in offering the Society’s heartiest congratulations to:
Dr. P. D. F. Murray on attaining his Doctorate in Science in the University
of Sydney.

Mr. A. F. Basset Hull on his election as a Corresponding Member of the
Academy of Natural Sciences of Philadelphia.

Dr. A. B. Walkom on his election as General Secretary of the Australasian
Association for the Advancement of Science in succession to Mr. E. C. Andrews.

Mr. E. C. Andrews on the invitation extended to him to deliver the Silliman
Lectures at Yale University in 1927. e

Mr. R. H. Cambage on his election as President of the Australian National
Research Council and as President-elect of the Australasian Association for the
Advancement of Science.

Dr. Margaret O’Dwyer on attaining the degree of Doctor of Philosophy of the
University of London.

During the past year the names of twelve members have been added to the
roll, and two names have been removed from it, five members have resigned and
we have lost five members by death. The number of ordinary members on the
roll is now 168. Death has continued to take heavy toll among members, the

iv. PRESIDENTIAL ADDRESS.

losses for the past year including Miss E. E. Chase and Messrs. H. E. Finckh,
J. J. Fletcher, Charles Hedley and Sir Hugh Dixson.

Eleanor Emily Chase, who died at Roseville on 17th May, 1926, was born in
Victoria and was educated at St. Catherine’s Church of England Girls’ School,
Waverley and the University of Sydney. She entered the University in 1914 and
obtained the degree of Bachelor of Science, with honours in Zoology and
Physiology, in 1917, gaining at the same time Professor Haswell’s prize in
Zoology. She was appointed demonstrator in Zoology on graduation, and in 1923
was appointed lecturer. During the five years that she was a member of the
University staff she won the warm regard of her colleagues and of a large
number of students by her conscientious devotion to her duties as well as by her
charming personality. Her opportunities for carrying out research were limited,
but she contributed one paper on “‘A new avian trematode” to our proceedings in
1920. She accompanied two important collecting expeditions during recent years,
one to Barrington Tops, N.S.W., and the other to the Capricorn Group, Q. She had
been a member of this Society since 1921 and was also a member of the Royal
Zoological Society of New South Wales. In addition she was a Vice-President of
the Sydney University Science Society, a member of the board of directors of
the Women’s Union at the University, and secretary to the committee of the
National Council of Women on sex education.

The sudden death, at such an early stage in her scientific career, of one who
had done promising zoological work and had endeared herself to those with whom
her work brought her into contact, is a distinct loss to biological science.

Hugh Dixson, who died at Colombo on 11th May, 1926, was born in George
Street, Sydney, on 29th January, 1841. He was educated at Timothy Cape’s
Academy at Paddington, and while still a youth he joined his father, who was a
tobacco manufacturer, the business later becoming probably the greatest of its
kind in Australia. He was deeply interested in church affairs and was a generous
benefactor to the Baptist Church in New South Wales, having, with his wife
Dame Emma Dixson, originated almost every existing trust fund connected with the
Baptist Union, including the Aged and Infirm Ministers’ Fund, to which he gave
£10,000. For some time he was president of the Young Men’s Christian Association
and was a liberal supporter of the Boy Scouts. He was knighted in 1921. His
general interest in science was manifested in his continuous membershio of
scientific societies; he had been a member of our Society since 1887 and of the
Australasian Association for the Advancement of Science since 1898. He was
particularly well known for his interest in amateur gardening, his pet hobby in
this respect being the growing of orchids.

Herman Edward Finckh, who died at Mosman on 31st May, 1926, was born
at Moore Park, Sydney, on 26th May, 1864. His earliest recollections were walks
on Sunday mornings, across what is now the Golf Links, where he used to watch
the aboriginals cooking over their camp fires. When ten years of age, he and
his younger brothers and sister were taken to Heilbron, Germany, to be educated;
he remained there for eight years, after which he toured Europe and America for
two years. On his return to Sydney he went into partnership with his father,
who carried on business as a jeweller in George Street, eventually taking over
the business himself. He retired about fifteen years ago and devoted himself
to natural history work. His earliest interest was horticulture and later on
the breeding of birds and small animals, in which he was very successful. In
1904 he decided to take up marine work—an interest that grew from a desire of

PRESIDENTIAL ADDRESS. Vv.

his daughter (now Mrs. C. A. Messimer) to have a salt water aquarium. He
began with one small tank, but later on when Mr. E. R. Waite resigned from the
Australian Museum, Finckh bought his tanks. From that time onwards his
tanks increased in number and he spent considerable time in studying the life
histories of the Australian fishes and also many exotic species that he imported
from time to time. His aquaria were well known to the naturalists of this State
and he was always ready to show them to anyone who was interested. He was a
recognized authority on the care and management of aquaria and during the
past two years was engaged upon a book dealing with these matters and also
on the life histories of some fish. It is hoped that this book will be published
shortly. At the time of his death his fish numbered about 5,000 mature specimens.

Finckh joined this Society in 1908; he had also been on the Council of the
Royal Zoological Society of New South Wales, of which he was a Vice-President
for many years; he did very efficient work as Hon. Treasurer of the Naturalists’
Society of New South Wales for many years; he was also a member of the
Aquarium Society, of the Wild Life Preservation Society and of the Gould League
of Bird Lovers. His keen interest in and love for nature made him a very
enjoyable companion and he never spared himself in his performance of his duties
in connection with Natural History in New South Wales.

Joseph James Fletcher was the eldest son of the late Rev. Joseph Horner
Fletcher. He was born in Auckland, N.Z. in 1850 and came to Queensland in 1860.
He was educated at Ipswich Grammar School, at Newington College (of which
his father was then Principal) and at the University of Sydney, gaining his B.A.
degree in 1870 and his M.A. degree in 1876. In 1870 he joined the senior staff
of Wesley College, Melbourne, and in 1876 went to England to study Biology at
the University of London, where he obtained the B.Sc. degree in 1879, being one
of the first two Australians to take a science degree from that University. Before
returning to Sydney he spent some time in study at the University of Cambridge
and there wrote his first paper in collaboration with J. J. Lister. This paper
was published in the Proceedings of the Zoological Society of London for 1881.
On his return to Sydney in 1881 he joined the staff of Newington College, where
he had as a colleague R. T. Baker, until lately Curater of the Technological
Museum, Sydney. He joined this Society in 1881 and was elected to the Council
on 31st January, 1883. He soon attracted the notice of its founder, Sir William
Macleay, who invited him to become its director and librarian. He entered upon
his duties on the first day of January, 1886, and from that time devoted all his
energies to the work of the Linnean Society of New South Wales. How he carried
out his life work can be judged by the prominent position this Society takes in
the scientific world. For 33 years he guided its affairs, edited its proceedings
and when, in March, 1919, he retired from the position of Secretary, he was
chosen as President, an office he worthily filled for two years. He remained on
the Council until his death at Hunter’s Hill on the 15th May, 1926. His member-
ship of the Society, with one exception, had been longer than that of any other
member of the Society.

The high degree of excellence reached in the performance of his duties is
known to all workers in natural science in Australia. The PRocrEDINGS since
1886 and up to the time of his retirement may justly be said to be a monument
to his capacity, his zeal and his devotion to duty. Authors of papers well know
that he was ever ready to advise them and if necessary to make suggestions, which
would tend towards the improvement of a paper in some detail and having set

Vi. PRESIDENTIAL ADDRESS.

a standard his watchful eye saw to it that such standard was maintained through-
out the series. In all his doings he not only commanded our respect, but our
affection as well and he served as an inspiration to many a student. Many of the
leading members of the Society at the present time owe a deep gratitude to him
for the valuable advice and instruction that he was always ready to give. Fletcher
had a profound reverence for the founder of this Society, Sir William Macleay,
and at all times instilled into our members the great debt.we owe to him. On the
14th June, 1920, to commemorate the one hundredth anniversary of the birth of
Sir William Macleay, he delivered an important address entitled “The Society’s
Heritage from the Macleays’’. In this he dealt with Alexander Macleay and his
two sons, William Sharp and George, and it furnishes a most valuable record
of what members of that family did in the cause of natural science. He had
almost completed the remaining portion dealing with Sir William Macleay.
Fletcher, no doubt, felt the removal of the office of the Society from Elizabeth
Bay to 16 College Street, but he did nothing to oppose this step, for he foresaw
that the new home nearer the centre of Sydney would increase the usefulness
of the Society.

His published papers include 36 in These ProceEepines together with one in
collaboration with A. G. Hamilton and another with C. T. Musson, who has also
presented for publication an important paper on Grevillea hybrids, based on the
careful notes made by Fletcher over a period of several years. This paper will
appear in 1927.

In January, 1900, Fletcher presided over the Biology Section at the Melbourne
Meeting of the Australasian Association for the Advancement of Science. In 1921
he was awarded the Clarke Memorial Medal by the Royal Society of New South
Wales. He was a trustee of the Australian Museum and a member of the Royal
Zoological Society of New South Wales.

After his death Mrs. Fletcher felt that the desk, presented to her husband
on his retirement trom the position of Secretary in 1919, should find a place in
the Society’s rooms, so she graciously presented it to the members. It is now in
the office and will be used by the future secretaries of the Society.

An appreciation of Mr. Fletcher by one of his oldest friends, Sir W. Baldwin
Spencer, appears at the conclusion of this part.

Charles Hedley, who died at his home, ‘“Nukulailai’, Muston Street, Mosman,
on the morning of the 14th September, 1926, was born at Masham Vicarage,
Yorkshire, on the 27th February, 1862. On account of his health his school life
was limited to two years at Eastbourne College, but he was an extensive reader
and received instruction from his distinguished father, Rev. Canon T. Hedley, a
Fellow of Trinity College, Cambridge. To avoid the English winters he was sent
each year either to the south of France or to Italy, and at these places he began
to find his first enjoyment of nature. At Mentone he purchased a book by A.
Moquin-Tandon on Conchology. This book on land shells adopted what was then
a modern view, that in the Mollusca the animal should be studied as well as the
shell it formed. He has often spoken of this book as the one that gave him a
stimulus which he carried throughout his zoological career. Here, when out
collecting, he met George French Angas, a former Secretary of the Australian
Museum, who was engaged in a similar occupation. Angas later gave him a
letter of introduction to Dr. G. Bennett of Sydney.

He left England for New Zealand in 1881, intending to learn sheep farming
there, but very soon found that the wet climate of New Zealand did not suit his

PRESIDENTIAL ADDRESS. Vii.

health. The conditions under which he worked were far from satisfactory, so he
came to Sydney in September, 1882. Shortly after his arrival he met Dr. G.
Bennett, to whom he was immensely drawn. He wrote to his father, “I have again
been to dinner with Dr. Bennett. Fancy speaking to a man who has talked with
Cuvier, who is an intimate friend of Owen, who knew or knows Darwin, Hooker,
Huxley, Lyell, Wallace, Thomson, Rolleston, Balfour, Gunther, all the distinguished
men of science who have lived in the last half-century!” A trip to Hay to find if
the dry inland air would relieve his asthma, was not successful, so he went to
Queensland. He lived first at Nerang and afterwards took up an oyster lease at
Stradbroke Is., Moreton Bay. Here, as in after years, he found he always had
better health when living close to the sea. Though he was able to visit the
Queensland Museum on many occasions he felt himself isolated from congenial
companionship. In 1884 he was fruit growing at Boyne Island, Port Curtis, where
he gathered information from the aboriginals for his first paper, “Uses of some
Queensland Plants” (Proc. Roy. Soc. Queensland, 1888). Here when driving in a
sulky he met with an accident, smashing his left elbow, and when he recovered
he found he could no longer do any laborious work and so he left Boyne Is. in
1888 and for about two years lived in Brisbane. He occupied himself in collecting
and doing voluntary work at the Queensland Museum. He found his first new
species, a slug at Burleigh Heads and described it as Limax queenslandicus with
figures and was pleased that his drawings caused such favourable comment. He
also published a list of the Land Shells recorded from Queensland.

On the ist January, 1889, he was appointed a supernumerary officer of the
Queensland Museum and on 2nd May, 1889, he became a Fellow of the Linnean
Society of London, and in July, Hon. Secretary of the Royal Society of Queens-
land. In 1890 at the invitation of the Administrator, Sir William Macgregor, he
visited British New Guinea, and reached some hitherto unexplored regions. He
spent some time at the St. Joseph’s River, Milne Bay, and made important
collections. Fever forced him to return to Brisbane though he wished to remain.
He came to Sydney towards the end of 1890 and resided there until his death.
On the 1st April, 1891, he commenced duty at the Australian Museum as assistant
in charge of Landshells. On ist January, 1896 he became conchologist and on
20th December, 1908, Assistant Curator. On the death of Robert Etheridge, Junr.,
in January, 1920, he was appointed Acting Director until his appointment on 14th
February as Principal Keeper of Collections. He resigned on 30th March, 1924, to
become Scientific Director to the Great Barrier Reef Committee.

Hedley’s connection with our Society began on the 25th June, 1890, when he
was elected a corresponding member. On 29th April, 1891, he became an ordinary
member; he served on the Council from 1897 to 1924 and was President, 1909-1911.
He contributed to this Society forty-four papers—the first in 1890—and five others
of which he was a joint author. These papers dealt chiefly with Mollusca, of
which he described numerous new species. His papers on Zoogeography,.a subject
in which he was keenly interested, are very important and he has made note-
worthy contributions to the study of the distribution of plants and animals in the
South Pacific. He also wrote a few papers on Ethnology. In all, he wrote nearly
two hundred papers which have been published in Australia and abroad.
Hedley’s papers were remarkable for the clear way he put his facts and for
the interest he excited in everything he wrote. His public lectures were always
crowded and, speaking in a simple way, with frequent humorous touches, he
always carried his audience with him.

Vill. PRESIDENTIAL ADDRESS.

He was an original member of the Australasian Association for the Advance-
ment of Science, was President of Section D at the Brisbane Meeting in 1909 and
had attended most of the meetings since its inception and had assisted the
Association in many ways. He had been President of the Royal Society of New
South Wales (1914), the Royal Zoological Society of New South Wales and the
New South Wales Naturalists’ Club. He was a member (1893) and Vice President
since 1923 of the Malacological Society of London, a corresponding member of the
Zoological Society of London (1922), a corresponding member of the Academy of
Natural Sciences of Philadelphia (1893), Honorary Member of the New Zealand
Institute (1924), Honorary Fellow of the Royal Society of South Australia and
corresponding member of the Royal Societies of Queensland and of Western
Australia. In 1916 he was awarded the David Syme Prize and in 1925 the Royal
Society of New South Wales presented him with the Clarke Memorial Medal in
recognition of his services to Australian Zoology.

In addition to his writings Hedley was a keen explorer. He was always
ready to join any expedition or trip to a place he had never visited before. He had
explored most of Eastern Australia. He had visited the Gulf of Carpentaria with
Dr. W. E. Roth, had led two trips to the Torres Strait Islands besides many smaller
ones in Queensland and New South Wales. He was deeply interested in deep sea
dredging and organized expeditions to explore the depths of the ocean. In
1905 and 1906 he was successful in obtaining a large collection of fish and
invertebrates from a depth of a mile from the ocean bed off Sydney Heads—a feat
never before attempted in Australian waters except from a man-of-war. With W.
L. May he was the first to explore the 100 fathom zone off the Tasmanian coast and
in company with Dr. J. C. Verco he lifted off Cape Borda the first deep sea bottom
of South Australia. He was one of the few scientists to put on a diving dress
and descend to a depth of 40 feet below the surface of Sydney Harbour. Of
later years his chief interest was the study of the Great Barrier Reef, and
his knowledge of it surpassed that of any other scientist. He had travelled over
most of it and in his many trips he had landed at a number of the islands of the
Reef area. He was the first civilian to make a trip by aeroplane over a portion
of the Reef. His presence on the Pan-Pacific Excursion to the Reef in 1923 added
greatly to the enjoyment and profit of those who were privileged to travel in the
S. S. “Relief” from Mackay.

Hedley did not confine his trips to Australia alone. He visited and collected in
New Guinea, Ellice Group, New Caledonia and from all of these places he obtained
valuable material. In 1896 he joined the expedition sent out by the Royal Society of
London under Professor Sollas to bore the atoll of Funafuti, Ellice Group. He
remained on the island for about 10 weeks and succeeded in amassing an interesting
collection, particularly of invertebrate and ethnological objects, together with much
valuable scientific information. An account of this work was given in “The Atoll of
Funafuti’ (Australian Museum Memoirs, iii, 1896-1900), to which Hedley
contributed the general account and the sections of ethnology and mollusca. Of
later years he desired to go further afield to see other parts of the world,
so in 1922 we find that he was in Western Canada and Alaska and in 1925 he
visited Africa to see the Great Rift Valley. In the last few years the two men
whose names will always be remembered in connection with Australian Zoology
have passed away—the late Professor W. A. Haswell and the late Charles Hedley.
Both in their own way have done more than any other two men to advance
Zoology in Australia. Haswell will always be remembered for his accuracy and

PRESIDENTIAL ADDRESS. ix.

care of detail and he has amongst zoologists a world-wide reputation for his
Text Book of Zoology. Hedley was known for his hospitality, being always the
first to welcome any visiting biologist, to make them feel at home in Australia,
and to place himself and his knowledge entirely at their disposal this all in
addition to the reputation he had for his scientific writings.

Hedley was held in the highest esteem by those who were his contemporaries
and he was an ideal companion during a walk through the bush or along the sea-
shore. He was of the utmost help to those younger than himself and many of
the members of this Society will remember with gratitude the advice that he was
ready to give them. He was always willing to listen to any biological or
geological problem that presented itself-and never grudged the time spent in
helping others in their work. By nature he was kindly and courteous and he was
the best of friends. Zoological Science in Australia has lost a great man, who
could ill be spared, but he has left behind him a monument of hard and patient
work and others will arise to carry on the work he has so ably begun.

The assistance he gave to young naturalists and to institutions has heen great.
During his lifetime he gave many books to this Society and to the Australian
Museum and by the wish of Mrs. Hedley, her husband’s books are being divided
between these two institutions, so our library will be increased by the addition of
many important books.

During the end of April until August, 1926, he had been supervising ths
sinking of the bore on Michaelmas Reef, Oyster Cay, near Cairns, and returned
to Sydney about the middle of August to make preparations to visit Japan in
connection with the Third Pan-Pacific Science Congress. He had to cancel his
arrangements as he did not feel too well, though no serious consequences were
expected and it was thought that a few weeks’ rest would restore him to health.
His sudden death on the morning of the 14th September, 1926, was a great shock
to all his friends. His remains were cremated and the ashes taken by Mr. E. C.
Andrews and in the presence of the other Australian delegates to the Third Pan-
Pacific Science Congress, distributed on the waters near the Great Barrier Reef,
which he loved and knew so well.

The year’s work of the Society’s research staff may be summarized thus:

Dr. R. Greig-Smith, Macleay Bacteriologist to the Society, has continued his
investigations into the activity of the mineral colloids such as fuller’s earth,
kaolin, silica, aluminium hydrate upon fermentation. The possible assistance given
to the alcoholic fermentation by their aiding the diffusion of carbon dioxide has
been reported upon when it was shown that the colloids have an action of their
own quite unconnected with the physical elimination of carbon dioxide. It has
been shown that agar-fibre acts like the mineral colloids in accelerating the
fermentation of solutions of sugar by yeast provided that a sufficient amount is
present. The possible stimulation of enfeebled yeast cells by the mineral colloids
led to the determination that when yeast cells are heated to just short of the
lethal temperature they have a different action upon saccharose than normal
unheated yeast. In presence of a mineral colloid the normal yeast inverts less
saccharose and generally consumes less sugar while the heated yeast inverts
the same amount of saccharose and consumes more sugar. It would seem that
heating the yeast partly diminishes its invertase-secreting function. During the
gum fermentation of saccharose by Bac. vulgatus it was noted that the non-
reducing sugars decreased to a minimum then increased and again diminished.
This led to the discovery of a substance intermediate between gum-levan and

x. PRESIDENTIAL ADDRESS.

levulose. Its presence can be shown but it has not been possible to prepare
it in the pure state or in pure solution; it is always accompanied by levulose.
Work is being prosecuted in the direction of the reason for the activity of the
mineral colloids in accelerating fermentation.

Miss May M. Williams, Linnean Macleay Fellow of the Society in Botany,
has continued her investigations into the Cytology and Phylogeny of the
Siphonaceous Algae, part 2 of which, dealing with Oogenesis and Spermatogenesis
in Vaucheria geminata was published in part 2 of the proceedings for 1926. She
has, further, been studying gametogenesis in Bryopsis plumosa with encouraging
results. The establishment of a marine aquarium in the laboratory has enabled
her to observe many interesting phases of gametogenesis. With the exception of
the formation of gametes by vacuolization instead of by cleavage, the details of
gametogenesis in Bryopsis plumosa are very similar to those in Codium tomen-
tosum. She is investigating the problem as to whether reduction in the number
of chromosomes in the nucleus occurs during gametogenesis, as in Codium
tomentosum, or at the first division of the zygospore following on fertilization.
As opportunity offers she is also continuing her search, as yet unsuccessful, for
the reproductive organs of Caulerpa.

In continuation of her work on the anatomy of certain ferns, Miss Williams
has completed ready for publication a paper on “The anatomy of Cheilanthes
vellea’, showing that this species exhibits, both in external morphology and
internal anatomy, well marked adaptations to its xerophilous habitat.

During the coming year she proposes to continue her investigations into
gametogenesis in Bryopsis plumosa and to undertake a critical examination of
the gametangia of Hctocarpus.

Dr. I. M. Mackerras, Linnean Macleay Fellow of the Society in Zoology, has
continued his investigations on the Diptera, having during the past year devoted
most of his attention to the Culicidae. The results of his work are to be
submitted in a series of papers under the title “Notes on Australian Mosquitoes”,
parts i and ii being ready for presentation. Part i, “The Anophelini of the
Mainland’, includes keys and figures which should make the recognition of the
males, females and larvae of all species occurring on the mainland relatively
easy. New life histories are described and notes are given on the status, habits
and distribution of the species. Part ii deals in a similar way with the mainland
and Tasmanian species of Ochlerotatus.

In addition to completing his work on the mosquitoes Dr. Mackerras proposes
to resume his studies of the comparative morphology and distribution of the
genera of the Tabanoidea and to proceed with the study of other families of
Diptera Brachycera.

Seven applications for Linnean Macleay Fellowships, 1927-28, were received
in response to the Council’s invitation of 29th September, 1926. I have pleasure
in reminding you that the Council re-appointed Miss May M. Williams and Dr.
I. M. Mackerras to Fellowships in Botany and Zoology respectively and appointed
Miss Ida Alison Brown and Miss Hazel Claire Weekes to Fellowships in Geology
and Zoology respectively for one year from 1st March, 1927. On behalf of the
Society I have pleasure in wishing them a very successful year’s research.

Miss Ida Alison Brown graduated in Science at the University of Sydney
in March, 1922, with First Class Honours and University Medal in Geology
and Second Class Honours in Mathematics, also being awarded the Deas-Thomson
Scholarship in Mineralogy and a Government Science Research Scholarship. The

PRESIDENTIAL ADDRESS. xi.

latter scholarship was, however, relinquished by her in order to take up the
position of Demonstrator in Geology, which she has held until her present appoint-
ment to a Linnean Macleay Fellowship. Since graduation she has carried on
research in geology and has published four papers, two in the Journal of the
Royal Society of New South Wales and two in our own proceedings. The first
of these papers, “Notes on hornblende and bytownite from hypersthene-gabbro,
Black Bluff, near Broken Hill’, is a petrological study, and the other three are
the result of field work in the districts of Milton and Moruya. For her work
as a Fellow Miss Brown proposes to continue her work on the geological history
of the South Coast district of New South Wales, studying in particular the
geological age, conditions of sedimentation, mutual relations and tectonic history
of the sedimentary rocks, as well as the relationships, petrogenesis and correlation
of the associated igneous intrusions, commencing in the Moruya district and
working south to Bodalla, Narooma, Mount Dromedary, Cobargo and Bega. Miss
Brown comes to us with the strongest recommendations from Professors David
and Cotton and she is the first woman to be appointed as a Fellow to do field
work in Geology. Though her proposed researches involve arduous work in
the field we may look forward to a satisfactory and successful year’s work.

Miss Hazel Claire Weekes, who graduated with First Ciass Honours in
Zoology at the University of Sydney in 1925, is the first woman to have been
awarded the University Medal in Zoology at graduation. She was also awarded
a Government Science Research Scholarship. In 1925 she commenced research
work on reptilian embryology and published, in conjunction with Professor
Harrison, a paper “On the occurrence of placentation in the Scincid lizard,
Lygosoma entrecasteauxvi’. This paper appeared in our proceedings for 1925.
These researches have been continued and a large amount of material has been
accumulated, two investigations, into the reproductive phenomena in Lygosoma
quoyi and the development of Amphibolurus barbatus being almost complete. This
line of work gives promise of extremely important results on the morphological
side of zoology and Miss Weekes proposes to continue her studies on reptilian
embryology during her tenure of a Fellowship, and we wish her a most
successful year’s research.

I wish to bring under your notice, more particularly those of you who are
entomologists, an important book written by a member of this Society, Dr. R. J.
Tillyard, M.A., F.R.S., and published in Sydney by Messrs. Angus and Robertson,
who have presented our Library with a copy. This book is entitled “The Insects
of Australia and New Zealand”. In this work, Dr. Tillyard opens with the classi-
fication of insects and gives a short account by which the 24 orders can be
recognized from one another. The derivations of the names of the orders and the
higher divisions are a help in understanding the various kinds of insects placed
therein. A census of the insects of the world, Australia and New Zealand shows
the large number of different species already known. More than one-third of
Australian insects belong to the order Coleoptera and in New Zealand slightly more
than one-half belong to the same order. Over 470,000 different insects have been
described from the world and of these 37,000 (about 8%) are found in Australia
and 8,000 (under 2%) in New Zealand. The number is being added to each year
and it will not be long before 50,000 insects are known from Australia. The orders
Plectoptera, Perlaria and Trichoptera are proportionately better represented in New
Zealand than in Australia. Chapter ii deals with a general account of the external
morphology, more detailed accounts being given in the chapters dealing with the

X11. PRESIDENTIAL ADDRESS.

various orders. In this chapter Dr. Tillyard explains the modern views on wing
venation and gives an instructive account of the latest system of nomenclature
and notation for the veins of the insect wing. Chapter iii gives a general outline
of the internal morphology of insects, giving a short account of the alimentary,
excretory, respiratory, circulatory, nervous and reproductive systems as well as the
organs of sense. In Chapter iv a concise account of the complete life cycle of an
insect is given under the headings: the egg or embryonic stage, the larva or
growing stage, the pupa or resting stage and the imago or sexually adult stage.
Chapters v-xxviii deal with the twenty-four orders in more detail, and treat of
the suborders, superfamilies, families and in some cases subfamilies mentioning
typical genera and species in each group. The whole is well illustrated with
details of structures, particularly wing venation and figures, both coloured and
black and white, of important species.

The subclass Apterygota contains three orders of insects which are entirely
wingless: (1) Thysanura (Bristle-tails, Silver-fish). In their early stages these are
very similar to the adult and very little work has been done on this order. Only
a few species are represented, but it is pointed out that many new forms will
probably be found. (2) Protura. These small insects have not yet been found
in Australia or New Zealand, but a short account of them is given so that they
may be recognized when found. (3) Collembola (Spring-tails). These small
and curious insects like the Thysanura and Protura are wingless, but they have
no very near relations amongst the insects. It has even been suggested by some
authors that they belong to a separate class distinct from Insecta.

The subclass Pterygota contains all the winged insects, though some few of
them do not possess wings. They are divided into two large groups. (A) Eeto-
pterygota in which the wings develop as external buds during the larval stages and
there is no true pupa. (4) Plectoptera (May-flies). Delicately formed insects,
usually small, always found near water, in which their early stages are passed.
The hindwings are often very small. About twenty species are known from
Australia and New Zealand. (5) Odonata (Dragonflies, Damselflies). ‘These
insects also as far as is known pass their early stages in water. They are
carnivorous and both nymphs and adults feed upon mosquitoes in their various
stages. Dr. Tillyard deals with this group very fully. (6) Orthoptera (Cock-
roaches, Mantids, Phasmids, Locusts, Grasshoppers, Crickets). In this order
the mouth parts are typically mandibulate, the larval forms closely resemble the
adults except for the absence of functional wings. The order includes five very
distinct superfamilies, which are considered as distinct suborders or even distinct
orders by some writers. The order contains some well known insects as cock-
roaches and grasshoppers and also some with very strange shapes. (7) Isoptera
(Termites, White Ants). These insects are more nearly related to cockroackes
than to the true ants. They are abundant in Australia especially in the north,
but only four species are found in Tasmania and three in New Zealand. They
cause many thousands of pounds loss each year by their attacks on houses, fences,
and railway sleepers. (8) Dermaptera (Harwigs). These are not very numerous
in Australia and only three species have been recorded from New Zealand where
a common species introduced from Europe is becoming a serious pest to fruit-
growers and gardeners. (9) Perlaria (Stone-flies). These insects form a small,
but well marked order about equally represented in Australia and New Zealand.
Their larvae are aquatic and are a valuable food for trout and freshwater fishes.
(10) Embiaria (Webspinners). A small order resembling Termites. Only six

PRESIDENTIAL ADDRESS. xiii.

species are known from Australia and none from New Zealand. (11) Zoraptera.
Only six species of this order have been described, none of which is found in
Australia and New Zealand. (12) Copeognatha (Psocids, Book-lice). Chiefly very
small insects, some of which are a pest in libraries and museums. (13) Anoplura
(Lice). Both biting and sucking lice are included in this order, the Mallophaga
(Biting lice, Bird lice) are represented by 88 species parasitic upon birds and
marsupials, whilst only four native species of Siphunculata (Sucking lice) are
known, though many introduced species occur. (14) Thysanoptera (Thrips).
Small elongate insects usually infesting blossoms. This order is well represented in
Australia about 14% of the known species occurring there. (15) Hemiptera
(Cicadas, Plant Hoppers, Scale Insects, Bugs). The insects of this order are
generally harmful, nearly all the Homoptera, and the majority of the Heteroptera
being plant feeders causing destruction to vegetabie foods. Some few Heteroptera
are beneficial, feeding on other insects. Scale Insects, Plant lice and other
Hemiptera are responsible for the greater part of the total damage done to our
crops and fruit.

The division (B) Endopterygota, to which the remaining orders
belong, have larvae in which the wings develop internally and have a true pupa or
resting stage. (16) Coleoptera (Beetles). This is the largest order of living
insects, comprising over 40% of the known insect fauna. They are an easily
recognized group, the forewing being specialized as a hard horny covering for the
hindwing. Many species are injurious owing to their attacks upon timber, [fruit
and grain. A few are useful as scavengers and others are predatory on scale
insects and aphids. The order is well represented in both Australia and New
Zealand. (17) Strepsiptera, a small order of minute insects parasitic upon the
orders Hymenoptera and Hemiptera (Homoptera). No species of the order have
so far been found in New Zealand. (18) Hymenoptera (Ants, Bees, and Wasps).
This is one of the most distinct of the Orders of Insects, usually with four stiff
membranous wings, though in some groups many of the species are wingiess
especially in the females. In many species the ovipositor of the female is often
specialized as a sting. Some few species may be considered injurious insects and
some ants and wasps are obnoxious, but generally the order may be looked upon
as very beneficial to man. Parasitic Hymenoptera play a great part in the main-
tenance of the balance of insect life, whereby many injurious species of other
insects are prevented from overruning the world. Ants are useful as scavengers.
(19) Neuroptera (Alder-flies, Lacewings). Most of the species of this order are
delicate looking insects. Many of the larval forms have prominent jaws. The
pretty antlions belong to this order, and also the delicate Psychopsis which are
often mistaken for moths. (20) Mecoptera (Scorpion-flies). A small order with
only twelve species in Australia and one in New Zealand. The perfect insects are
diurnal, mostly inhabiting damp cool places. Some are predaceous, killing many
small insects. (21) Diptera (Two-winged flies). In this order, except for ihose
species that are wingless, the forewings are only developed, the hindwings being
reduced to a pair of small club-like appendages called halteres or balancers. ‘rom
New Zealand almost as many species have been described as from Australia, but
this is chiefly due to the small amount of attention this group has received in the
latter country until very recently. In the last two or three years many new
species have been made known from Australia and though the number of flies,
given by Dr. Tillyard as 2,120 when he wrote his book, is only about 4% of the
world’s total of flies, if a census were now taken the percentage would be nearer

Xiv. PRESIDENTIAL ADDRESS.

six. It is probable that not half the flies have yet been described from Australia.
Flies are undoubtedly the most important order of Insects considered in relation
to man. They are discussed from their medical aspect in another portion of this
Address. Besides their importance as carriers of disease, many species cause great
destruction to trees, fruit and vegetables. Some few flies are certainly beneficial;
the robber-flies prey on other insects. The larvae of Hover-flies prey on aphids
and larvae of other groups are parasitic upon the larvae of other insects.
(22) Siphonaptera (Fleas). This order is highly speciailzed and wingless. They
cause irritation by their biting and are carriers of disease. (23) Trichoptera
(Caddis-flies). This order is related to the Lepidoptera, in fact they are often
mistaken for moths, but their wings are covered with dense hairs and rarely with
seales. The early stages are passed entirely in water, the larvae living in portable
cases or fixed abodes, often very curiously shaped. The larvae form one of the
important articles of diet for freshwater fishes. In New Zealand the order has
been more extensively studied than in Australia, from the latter country many
more than the sixty species already described will be made known. (24)
Lepidoptera (Butterflies and Moths). This is the best known order of insects on
account of the striking beauty of many of the adult forms. The perfect insects do
little damage, and are useful in the fertilization of flowers, but the larvae cause
an immense amount of destruction.

Chapters v to xxviii are all designed on the same plan, and consist of sections
dealing with Morphology, Life History, Distribution, Fossil History, Economics
and Classification. The illustrations are copious, clear and in most cases original.
Each chapter concludes with references to the more important papers to be
consulted for further details. Dr. Tillyard gratefully acknowledges the help he
has received from various entomologists in the various orders in which they
specialized.

The chapter on the fossil record and origin of the Australian and New
Zealand Insect Faunas shows us what has been accomplished in the space of a
few years. Before Dr. Tillyard began to study fossil insects very few species had
been recorded, but his enthusiasm inspired several collectors, with the result
that a large number of specimens had been found and described. The number is
still increasing and Dr. Tillyard has already received a large number of new
specimens to study.

The concluding chapter is devoted to the collection, preservation and study
of insects and contains very valuable information. A glossary of technical terms
is provided as an appendix.

The chief importance of this valuable book to us is that it deals entirely with
the insects of Australia and New Zealand. In the vast amount of original matter
it contains it has a world-wide importance. Very few general entomological works
are aS comprehensive and still fewer as up to date. The modern lines of
research are included up to the date of writing. This is the second general
entomological work produced by a member of this Society, Mr. W. W. Froggatt’s
excellent “Australian Insects’ having been out of print for several years.

Our congratulations are due to all concerned in the production of this book,
to author, to artists, to printer and to publishers.

PRESIDENTIAL ADDRESS. KV.

MEDICAL AND VETERINARY ENTOMOLOGY IN AUSTRALIA.—A REVIEW.

In the address from this chair last year the retiring President, Mr. H. J.
Carter, gave a résumé of the evolution and development of entomology from early
times to the present day. In his address Mr. Carter omitted any consideration of
one important economic line of modern development—the relation of insects to
the carriage of disease both in man and animals. This branch of entomology was
deliberately omitted in order that I might deal with it in fuller detail than would
be possible in Mr. Carter’s more general address. It would be, of course, manifestly
impossible to traverse adequately the whole of this subject in the time at our
disposal tonight. It is my purpose to restrict my remarks mainly to advances
made by Australians in this particular field and at the same time to give a rapid
survey of the special problems that confront Australian workers in the lines of
medical and veterinary entomology.

The whole science of medical entomology is of quite recent development and
may be said to take its origin from the discovery by the late Sir Patrick Manson
in 1878 of the role played by Culex fatigans in the transmission of Filaria
bancrofti. This discovery established the principle of development of a metazoan
parasite in an insect host. The existence of a corresponding, in this case sexual,
development of a protozoan parasite in an insect host was shown by Ross’s
discovery in 1898 of the life history of the parasite of malaria. Ross was unable
from the lack of material to follow out all the stages in the malarial parasite, but
was able to do so in the case of the allied Proteosoma of sparrows. Since that
time it has been shown that a considerable number of parasites both of man
and animals pass portion of their life cycle—generally the sexual cycle—in the
bodies of insects.

Another aspect of medical entomology later came into prominence when it
was shown that bacterial agents of disease could be conveyed through the medium
of insects, particularly of flies and fleas. The transmission in this case is purely
a mechanical one and there is no question of the cyclical development of a
parasite.

The discoveries made by the protozoologist, pathologist and sanitarian in the
mode of transmission of insect-borne diseases naturally led to further study of the
insects themselves. Ross’s discovery of the vector of malaria and the American
Commission’s discovery of the transmission of yellow fever by Aedes (Stegomyia)
aegypti undoubtedly stimulated study of the mosquitoes as a whole and greatly
enhanced our knowledge of the group. Similarly, the discovery of the role played
by the common house fly—Musca domestica—in the transmission of typhoid,
dysentery and similar diseases led to more intensive study of the Diptera as a
whole, particularly of the house frequenting species. The discovery that trench
fever as well as typhus and European relapsing fever was spread by the common
body louse considerably increased our knowledge of the habits and bionomics of
this insect. In similar fashion the study of fleas followed on the revelation of
their spread of pestis bubonica and of ticks on the part played by them in the
transmission of piroplasmosis and allied diseases in cattle and other animals,
though consideration of ticks and mites and their relation to disease is rather
outside the scope of the present address.

Examples could be multiplied, but enough have been quoted to show the
necessity for systematic study and classification of the insect vectors both actual
and potential. I propose to discuss the advances made in Australia in the
systematic study of certain groups of Australian insects which are, or which may

XVi. PRESIDENTIAL ADDRESS.

contain, species of economic importance from the medical and veterinary aspect
as well as to outline the advances made in the study of the diseases concerned,
particularly from the point of view of their relations to their insect hosts. I have
excluded from my review ali consideration of the Arachnida and diseases borne by
this Class, though such consideration is generally nee aEd ee as falling within the
province of the medical entomologist.

Viewing the Class Insecta as a whole it will be seen that only certain groups
of insects are concerned in the transmission of disease, and that these are such as
by their habits of life history come into more or less close association with man or
animals. Practically all insects of medical or veterinary importance fall into
the orders Diptera, Siphonaptera, Hemiptera and Anoplura.

Diptera: Within recent years considerable activity has been shown in the
study of Australian Diptera, hitherto one of the more neglected orders of
Australian insects. Harly describers of Australian flies included Desvoidy,
Macquart, Walker, Schiner, Thomson and Bigot. Unfortunately, many of these
early descriptions are worthless and the species practically unidentifiable except in
cases where the types are still available for comparison.

From 1888 to 1890 F. A. Skuse contributed a series of eight papers with two
supplements to These ProcEEDINGS on the Diptera of Australia. His work was,
however, confined to the Nematocera, with the exception of two small papers on
Acalyptrate flies of economic importance. The only groups of medical interest
dealt with are the Culicidae or mosquitoes and a few blood-sucking midges
comprised in the families Ceratopogonidae ana Simuliidae.

Between 1914 and 1918 Arthur White published a series of papers on
Brachycera Orthorrhapha dealing mainly with Tasmanian species, but including
also papers on mainland species. His writings included descriptions of new
Tasmanian Tabanidae.

Since 1920 numerous papers on Australian Diptera have appeared, mainly
from the pens of Hardy, Malloch, Bezzi, Mackerras, Hill, Ferguson, Alexander
and Tonnoir. :

Culicidae: Skuse (1888) first studied the mosquitoes systematically and
described 19 new species. This was in continuation of his work on Australian
Diptera and was not undertaken from the medical standpoint.

The greater interest shown in the systematic investigation of mosquitoes
following on Ross’s discovery of the transmission of malaria, and in which such
men as Giles, Blanchard, Theobald, Dyar and Knab, Edwards, Christophers, ete.,
played a leading part, commenced in the ’nineties and has been continued to the
present day. In this general study of the group, Australian species, of course,
were included, mainly in the works of Theobald. In this connection full credit
must be given to the unremitting efforts of Dr. T. L. Bancroft of Burpengary and
later of Hidsvold to secure, either by capture or by breeding, representative
collections of mosquitoes from the districts in which he was resident. In 1908
Dr. Bancroft published a census of Queensland mosquitoes in which some 32 species
are listed. Though this comprised only about one-third of the species now known
the work was the only list available for many years and proved invaluable to
Australian workers.

The next advance in the systematic study of Australian Culicidae followed on
the establishment of the Australian Institute of Tropical Medicine in Townsville
and the appointment of an entomologist to that institution. During the years
1912 to 1920 Mr. F. H. Taylor, the entomologist to the Institution, published a

PRESIDENTIAL ADDRESS. XVil

series of papers in which a large number of new species of mosquitoes are
described. These included not only species from Australia, but also from New
Guinea.

About this period the scale classification introduced by Theobald broke down
by its own weight and was replaced by the more rational structural system first
proposed by the Americans and now adopted by Edwards of the British Museum.
The resultant changes in generic appellations caused a period of some confusion
in the nomenclature of Australian mosquitoes, which was increased by the rather
considerable specific synonymy that existed. Through the joint efforts of Mr. F.
W. Edwards, of the British Museum, and of Mr. G. F. Hill, then entomologist to the
Tropical Institute at Townsville, the confusion was finally cleared up and the
issue in 1924 by Edwards of his synonymic list of adult mosquitoes of the
Australasian Region has placed in our hands an authentic list for the use of all
future workers in this group.

Larval characters: While the adult Australian Culicidae are now fairly well
known much yet remains to be done on the identification of the larvae. Descriptions
of most of the common species have been published by Cooling, Taylor and Hill,
while in some instances the larvae of extra-limital species have been described
by workers outside Australia.

The latest worker in this field—Dr. I. M. Mackerras, Linnean Macleay Fellow
in Zoology—has already a paper on the Anophelini ready for publication and a
second on the Culicini nearly completed.

Fieldwork: Work on Australian Culicidae has not been confined to the
systematic description of new species. The work of Dr. T. L. Bancroft in
collecting and breeding out species in Southern Queensland has already been
mentioned. Similar work was carried out in New South Wales by Dr. J. B.
Cleland largely in connection with the animal experimental station on Milson
Island, Hawkesbury River. In his numerous journeys about the State Dr. Cleland
also carefully collected and recorded the species of mosquitoes encountered and
encouraged his officers to do likewise, with the result that it is now possible to
map out the distribution of the Culicidae in New South Wales with some degree
of accuracy.

In North Australia much of the early work of collecting was carried out
by G. F. Hill, particularly during his residence in the Northern Territory and
later in Queensland and New Guinea. In Northern Queensland Taylor, Priestley,
Breinl and Cooling all carried out extensive collecting. Taylor was responsible for
a series of mosquito surveys of Northern Queensland towns and also undertook a
survey of the mosquito population of the irrigation areas on the Murray River in
Victoria. Recently, series of mosquito surveys have been made of Queensland
towns mainly by Dr. Hamlyn Harris. Surveys have been made at various times
in and around several of the capital cities, notably Brisbane and Adelaide.

Mosquito eradication work: Only brief reference can be made to campaigns
that have been instituted for the eradication of mosquitoes in various parts of
Australia. The most complete and the best sustained appear to be the measures
instituted by the Health Department in Brisbane, which were initiated by Dr.
Elkington in 1912. Other centres in which anti-mosquito measures have been
put into operation include Adelaide, Toowoomba, Kyogle and various suburban
municipalities around Sydney.

In New South Wales powers are given under Ordinance 41 of the Local
Government Act, 1906, to enable local authorities to compel householders to

LB

xviii. PRESIDENTIAL ADDRESS.

prevent the breeding of mosquitoes in or around their properties. This Ordinance
is not in general use, but can be applied to any Municipality or Shire by
proclamation. The Ordinance compels householders to keep cisterns, tanks, etc.,
properly screened or covered, to empty any receptacles that may contain water,
to rectify defective water spouting, etc., to oil or stock with fish permanent and
ornamental collections of water and generally to take such measures as may be
necessary for the prevention of mosquito breeding. The more extensive work of
ditching, draining, etc., must, of course, be undertaken by the local authority in
each district. The stumbling block to the thorough carrying out of anti-mosquito
measures is always the financial one and though in many cases something is done,
the result often falls far short of expectations and it is often difficult to ensure
that adequate maintenance work is continued.

It is to be pointed out here that, whereas in tropical and sub-tropical Australia
the eradication, or at least control, of the mosquito is a question of public health
importance, in the more temperate portions of the continent this is not the case
and it is merely a question of abating a nuisance.

Mosquito-borne Diseases: Three distinct mosquito-borne diseases occur in
Australia—Malaria, Dengue Fever and Filariasis. Malaria was at one time
undoubtedly much more prevalent in tropicai Australia than it is to-day and while
endemic foci probably still exist, many of the outbreaks recorded of recent years
have been due to the reintroduction of the disease during the rainy season. In
southern Australia only sporadic cases have occurred, some half-dozen or so being
on record. (New South Wales 7, Victoria 1, Western Australia 1). While, however,
the disease is a relatively rare one even in the tropical belt of Australia, the vector,
the Anopheline mosquito, is widespread and has been recorded by various observers
(Hill, Taylor, Ferguson, Cleland, etc.) from almost every portion of the continent.
Much uncertainty exists as to the exact species of Anopheles that is responsible.
The common and widespread species, A. annulipes, occurs throughout the
temperate region, but appears to be, if not replaced by, at least associated with, a
closely allied species, A. amictus, in northern Australia. Do the comparatively
slight morphological differences between these two indicate, as Hill suggests, a
possible difference in their readiness to act as vectors for the plasmodium? Or is
there any difference in the avidity with which they will attack man? Reference
might be made in this connection to the researches of Roubaud and others into
the man or cattle attacking races of A. maculipennis of southern Europe and their
relation to the incidence of malaria. That A. annulipes can carry malaria is
shown by the sporadic cases in New South Wales. There are, of course, other
species in tropical Australia, notably A. bancrofti and A. punctulatus, which may
be responsible for the conveyance of malaria in these regions. It is to be noted
that in Australia no species has been actually proved to be a vector by experimental
feeding on malarial patients and afterwards finding the developing sporozoites on
dissection of the mosquito, though Heydon in 1923 proved by experiment that
A. punctulatus was an efficient carrier at Rabaul. Clearly the factors governing
the incidence of malaria, even in the tropical portions of Australia, are not fully
known and require investigation. The fact of Australia’s comparative immunity
may depend on questions of sparseness of population or, as mentioned above, on
relative unsuitability of the insect vector.

Dengue Fever: It is now definitely established that dengue fever is conveyed
through the medium of infected mosquitoes; the great part played by Australian
workers in the elucidation of the aetiology of the disease was the proof of a

PRESIDENTIAL ADDRESS. . xix.

particular mosquito—Aedes (Stegomyia) aegypti—-as vector. Graham working in
Syria first definitely proved that dengue was conveyed by mosquitoes and, though
his results appeared to incriminate Culex fatigans, it was admitted that his
stocks of this species probably contained individuals of Ae. (Steg.) aegypti.
Experiments by Ashburn and Craig were not more conclusive. In 1906 Dr. T. L.
Bancroft of Queensland definitely succeeded in transferring dengue by means of
Ae. (Steg.) aegypti in two instances. This positive result was overlooked for some
years and left some doubt as to whether other methods of infection could be
absolutely excluded since Bancroft was working in a dengue district. In 1916
Drs. Cleland, Bradley and McDonald of Sydney, in a series of well planned
experiments, definitely proved Ae. aegypti to be the vector. These observers
brought down Ae. aegypti from the northern rivers of New South Wales to
Sydney, where this species does not occur, and by feeding them on volunteers
succeeded in reproducing the disease in four instances, the diagnosis being
confirmed by inoculation experiments in other volunteers. Their results have
been confirmed since by workers in other parts of the world, notably in the
* Philippines. The Philippine Commission found that Ae. aegypti did not become
infective until eleven days after feeding on an infected patient. This is against
the whole evidence of the epidemiology of the disease, which indicates that the
mosquito may be infective during the first three days after feeding on a patient.
This was shown furthermore to be the case experimentally by Chandler and Rice
in Texas. It appears from a study of the Philippine Commission’s report that no
experiments were carried out with infected mosquitoes under seven days after
being infected and it may well be that there are two methods of infection—a
mechanical method of transference of infected material within the first three
days of infection of the mosquito and a developmental method in which the
parasite undergoes some change in the mosquito, which does not become infective
for eleven days. Further work is required on these points. The causal organism
of dengue itself yet remains to be discovered.

Dengue is endemic in northern Australia and invades southern Queensland
and northern New South Wales at irregular periods. When the disease makes its
appearance up to 90% of the population of a town may be infected. Possibly
the periodicity of the appearance of the disease may be associated with mass
immunity of the population from a previous epidemic. The fact that the vector
disappears from a locality during the colder months in the more temperate portions
of its habitat, passing the winter in the egg stage, accounts for the disappearance
of the disease with the onset of cool weather. Until the 1926 epidemic in New
South Wales dengue had only been recorded from towns along the north coastal
rivers, but in 1926 towns along the north-western slopes and on the north-western
plains were heavily infected. Enquiry showed the presence of Ae. aegypti
wherever dengue was present and revealed the fact that this species of mosquito
was much more widely spread inland than had been previously known. Whether
the inland spread was a recent one or whether the presence of the mosquito had
been merely overlooked could not be determined.

Filariasis: The third great mosquito-borne disease—filariasis—is widespread,
at any rate in certain of the Queensland coastal districts. The discovery of the
parent worm—Filaria bancrofti—was made by the late Dr. Joseph Bancroft of
Brisbane in 1876. The fact that the mosquito was the vector was discovered by
Sir Patrick Manson at Amoy, in China, in 1878, but the full details were not
worked out; this was largely done by Dr. T. L. Bancroft in Brisbane in 1899.

Xi. PRESIDENTIAL ADDRESS.

Filaria has been proved to be carried by a number of different mosquitoes, the
commonest being the night feeding mosquito, C. fatigans, and it was with this
species that Dr. T. L. Bancroft successfully worked in Queensland. Further work
by Miss M. J. Walker in Brisbane in 1923-4 confirmed Dr. Bancroft’s results with
C. fatigans. Both observers obtained partial development in C. annulirostris;
other species of mosquitoes yielded negative results. In the Pacific Islands the
common vector as shown by Dr. Manson Bahr (1912), is Ae. (Steg.) variegatus,
but this species does not touch Australia; furthermore, the island disease is non-
periodic and is associated with numerous cases of HElephantiasis, whereas the
Queensland disease is definitely nocturnal in periodicity and Elephantiasis is
extremely rare, while lymph varix, chyluria, etc., are not uncommon. A consider-
able field, therefore, yet awaits the investigator in Queensland into the aetiology
of the disease, work in which both the pathologist and the entomologist might be
suitably employed.

Tabanidae: This family, familiarly known in Australia as March Flies, are
abundantly represented, close on 300 species being known. Many of the better
known forms were described by Macquart and Walker in the early part of last
century. Ricardo, in a series of papers in the Annals and Magazine of Natural
History from 1900 to 1917, reviewed many of the older described species as well
as described many others as new. Australian workers were thus able to place
correctly many species whose identity would otherwise have been uncertain.
F. H. Taylor (1913-1920), of the Australian Institute of Tropical Medicine,
Townsville, in a series of six papers, four of which were published in These
PROCEEDINGS, added numerous species, mainly from northern Australia.

During the years 1919-20 additional species were added by Ferguson, Ferguson
and Henry, and Ferguson and Hill. These authors also discussed the somewhat
extensive synonymy that had grown up in the family.

Studies on the life histories of Australian Tabanidae were published by
T. H. Johnston and M. J. Bancroft and by G. F. Hill, but the life history of only
some half-dozen species is as yet known. In connection with the work on
Onchocerciasis extensive collections of Tabanidae have been made in various parts
of Australia in response to appeals made by the various committees appointed to
investigate this disease. These collections have led to a great increase in our
knowledge of the distribution of these insects as well as to the access of new
species.

Diseases conveyed by Tabanidae: While no disease has been definitely proved
to be conveyed by Tabanidae in Australia, strong suspicion falls on these flies as
the probable vectors of Onchocerca gibsoni, the causal organism of bovine
Onchocerciasis. This nematode disease is widespread in the Northern Territory,
Queensland and New South Wales and is responsible for severe economic loss.
While the causal organism is known, the mode of transmission is as yet unascer-
tained, though various facts point to the vector being a flying insect. HExtensive
feeding and other experiments with Tabanidae carried out at Kendall yielded
negative results, which could not be regarded as conclusive. Dissection in a few
cases showed the presence of larval filariae in the proboscis of Tabanidae. Similar
larvae had been found by Johnston and Bancroft in Queensland and named
Agamofilaria tabanicola. These authors regarded the larvae as probably immature
forms of some species parasitic in a native animal or bird. Reference might be
made here to certain suggestive experiments recorded by Dr. Cilento of Townsville.
In the one observation, Tabanidae were seen to bite two cattle on the brisket; the

PRESIDENTIAL ADDRESS. XX

spots were marked and later worm nodules found to have developed there. In the
second case the crushed up heads of several Tabanids, in which nematodes were
seen in the proboscis, were injected into the brisket and were followed by the
development of worm nodules. The value of these experiments was to a certain
extent impaired by the fact that they were carried out in a district in which the
disease is naturally common and other modes of infection could not be excluded.

The whole question of the transmission of Onchocerciasis was further com-
plicated by the discovery that two other species of Onchocerca—O. gutterosa and
O. lienalis—occur in Australian cattle and in no case is the vector known.

There is no evidence that Tabanids have ever been associated with the
spread of anthrax in Australia, as appears to have been the case in other
countries.

Other blood-sucking Diptera: The Leptidae, though usually non-blood-sucking
flies predaceous on other insects, contain a single Australian genus of blood-
sucking species, Spaniopsis, which is confined to this country. Some five or six
species are known, but they are of academic interest only, as they are not known
to be associated with any disease.

Simuliidae or Buffalo gnats are represented in Australia by fourteen described
and a number of undescribed species. The family was revised in 1925 by Tonnoir.

Ceratopogonidae: Minute midges belonging to the genera Culicoides and
Leptoconops are widespread in Australia and are commonly termed sandflies.
They are vicious “biters’, raising irritating papules which are generally more
irritant after twenty-four hours or longer. The various species found in
Australia have been by no means thoroughly investigated, though some half-dozen
have been described.

Psychodidae: A single species of Phlebotomus has been described from
northern Queensland by G. F. Hill. It is not Known whether it is associated with
any disease, but short duration fevers analogous to three day or Phlebotomus
fever are known to occur in the tropical regions of Australia.

Muscidae: Two blood-sucking species of Muscidae occur in Australia, both
introduced. Stomoxzys calcitrans—the stable fly—is widespread throughout the
continent and, though usually confining its attention to cattle and horses, will on
occasion attack man. It has been suspected as a possible vector of Onchocerca
gibsoni, but extensive experiments with calves failed to prove any causal relation.
S. calcitrans has been accused, at various times, of transmitting infantile paralysis,
anthrax and surra, but there is no evidence that this species has ever acted as
vector of these diseases in Australia. ‘The stable fly is, however, known to be
the intermediate host of Habronema microstoma, a nematode parasite of horses
in Australia. The life history of this parasite and its allies—H. muscae and
H. megastoma—have been worked out by Hill, Bull, and Johnston and Bancroft.
Haematobia exigua (Lyperosia exigua), a close ally of Stomoxys calcitrans, has
been introduced into northern Australia and has spread from the Northern
Territory into the Kimberley district of Western Australia. The attacks of this
insect cause considerable irritation and the formation of intractable sores, leading
to marked loss of condition in the cattle attacked. The presence of the fly is
thus of serious economic importance and is the subject of special investigation by
the Commonwealth Government.

Non-blood-sucking Diptera: The vast number of Diptera are non-blood-sucking
and only a few are of interest as actual or potential carriers of disease. Certain
flies by habits of breeding or by predilection for certain food are attracted to

Xxil. PRESIDENTIAL ADDRESS.

human dwellings and come into close contact with man. Species that breed in
excreta and that are attracted to food stuffs are especially dangerous: Practically
all the important species are contained in the families Muscidae, Calliphoridae
and Sarcophagidae.

Muscidae: Musca domestica, the commor house fly, is par excellence the
species responsible for the spread of disease, particularly such diseases as typhoid,
infantile diarrhoea and dysentery. Since the discovery of the role of this species
in spreading typhoid fever, a discovery that followed on the Commissions of
Enquiry into the typhoid of the Spanish-American and South African Wars a vast
amount of evidence has accumulated incriminating this species and an extensive
literature has been compiled of its habits, bionomics, etc. While we thus have a
considerable mass of information on these points most of the evidence
has been obtained by study of the species in other regions and mainly in
temperate climates. Further information is required as to the bionomics of
M. domestica under Australian conditions, though in essentials the life history
will be found to be the same in all countries. There are, however, questions of
possible aestivation in the height of summer—it is a well known fact that flies
are less abundant in the summer than in spring or autumn; questions of the
relative abundance in different localities and States, to mention only two of the
many obscure but important aspects of the bionomics of the house fly under
Australian conditions. Such questions may be of paramount importance in the
study of disease. Why is infantile diarrhoea more common in a hot, dry summer?
Is this related to an increase in the fly prevalence during such seasons? What is
the relation of its local incidence to that of house flies? It is freely stated that
the seasonal curve of this disease closely follows that of the prevalence of flies.
As a matter of fact, there is no adequate data on which the latter curve could be
constructed for any part of Australia. To answer these and similar questions fly
counts spread over a number of years are necessary. Such fly. counts are
unfortunately not available, with the exception of a short series made by Dr. J. B.
Cleland some years ago in Sydney; their institution and maintenance over a series
of years is becoming a matter of urgent importance.

Prevention of Fly Breeding: Though the principles of the prevention of fly
breeding are well known, but little has been done to minimize the evil. Adequate
powers exist under various by-laws and local government ordinances to regulate
the storage and disposal of manure, the disposal of garbage and rubbish, the care
of stables, dairies, the construction and care of privies, etc., but though these
regulations may be, and generally are, enforced, many exceptions exist. I have
personally seen conditions existent in country towns which were a disgrace to the
community. The control of these conditions is, of course, a matter for the
sanitarian. The entomologist has done his share in pointing out the conditions
regulating fly breeding; he cannot do more. Too often, however, the efforts of the
municipal sanitary or health inspector are nullified by the apathy or even active
opposition of the local council. Better results might be obtained if all health
inspectors were directly under the control of the central health authority and not
servants of local councils. Conditions are as a rule better in the larger towns
and cities and the advent of the motor has helped to diminish the fly prevalence
by the decrease in the number of horses and the consequent lessening of the use
of stables in the cities. In some degree the fly prevalence in a community may be
measured by the typhoid incidence. Certain towns, particularly where water
carriage of sewage exists, are always free from the disease. This does not

PRESIDENTIAL ADDRESS. Xxili.

necessarily mean a lessened fly prevalence, but that the fly has not the opportunity
to come in contact with infected material. In other towns typhoid recurs annually.
That typhoid fever is diminishing with yearly fluctuations in New South Wales
is shown by the figures for the yearly incidence for the last ten years. To obtain
a proper indication of the spread of typhoid by flies it would be necessary to
deduct cases due to other causes, such as direct contamination of food, milk, etc.,
at the hands of typhoid carriers.

TABLE I.—TyPHOID FEVER.

Year. | Population. Cases. Deaths.
ee eee — —— —,
1917 1,886,701 1,091 103
1918 1,928,174 | 810 112
1919 2,000,173 857 | 106
1920 2,099,763 1,016 | 132
1921 2,128,786 | 949 | 129
1922 2,174,688 | 706 99
1923 2,211,106 873 104
1924 2,256,649 | 768 97
1925 2,306,081 | 533 80
1926 2,349,000 674 | —
|

Several other species of Muscidae closely related to the house fly have been
described from Australia; the nomenclature of these is by no means settled,
since some extend beyond Australian limits. The most important is, however,
Vivaparomusca ventustissima, the well known bush fly. This species is widely
distributed throughout Australia, and though it does not enter houses it will
persistently attack man in the open, being especially attracted by moist surfaces,
such as the eyes, nostrils, lips, or by open sores. Its prevalence varies greatly in
different years and there is reason to believe that the species is associated with the
spread of ophthalmic infections, such as conjunctivitis (“Sandy blight”) and
trachoma. It may also be the transmitting agent in cases of Bung-eye. This
species was extraordinarily abundant in the summer of 1923-24 and according to
the evidence of the school medical services this season was associated with an
unprecedented number of cases of ophthalmia in country schools.

An account of the bionomics of this species in Queensland was published by
Johnston and Bancroft.

Calliphoridae and Sarcophagidae: These two families may be briefly considered
together since in both cases the habits are similar. They are largely of importance
because of their habit of attacking meat and depositing their larvae and eggs
thereon, thus rendering it unfit for human consumption. The more severe
economic loss occasioned through the blowing of sheep will be considered under
myiasis. Both families are strongly represented in Australia. The Sarcophagidae
have been studied by Johnston and Tiegs and by Johnston and Hardy and in a
series of papers by these authors some 25 species are recorded from Australia.
The Calliphoridae have until lately been in a state of some confusion as far as
the nomenclature is concerned, despite attempts by Patton and by Hardy to
elucidate the family. I have recently received, however, a paper by Malloch on

XXiv. PRESIDENTIAL ADDRESS.

the family, which will be presented to this Society during the current year and
which will place the Australian species on a sound classificatory basis. The
question of the correct nomenclature is of importance, as Malloch found that under
several of the commonly accepted names two or more species have been included.
Differences of chaetotaxy and of male genitalia have been largely neglected in the
past, but these may be of great importance in determining specific status.

The bionomics of the Sarcophagidae have been largely studied by Johnston
and Tiegs and by Johnston and Hardy, and the latter author has also devoted
considerable time to the study of the bionomics of the Calliphoridae in Queensland.
Bionomic studies of the Calliphoridae were also made by J. L. Froggatt in his
investigations into the sheep-blowfly problem. Reference may here be made to
the series of experiments carried out by W. B. Gurney into the range of flight
of Australian blowflies. This observer found that marked blowflies could be
captured up to ten miles from their point of liberation.

Myiasis: By myiasis is understood the development in man or animals of the
larval and pupal stages of dipterous parasites. Such myiasis may be more or
less accidental or may be, as in the case of the bot flies, a constant phase in the
life history of the particular species of fly. Both the common species of blowfly—
Calliphora stygia and C. augur—may on occasion give rise to accidental myiasis,
generally in the nature of the blowing of open wounds. Cases of intestinal
myiasis are also known, the fly concerned being Fannia canicularis, a common
introduced species.

Much more important than these cases of human myiasis is the condition of
myiasis in sheep due to infestation mainly of the crutch with one or more of the
common blow or blue bottle flies. The condition has become so serious during
recent years that considerable economic loss has been occasioned, not only
through death of affected animals, but also through deterioration of the wool
through constant yarding, etc., for the purpose of jetting and crutching or other
remedial measures. In New South Wales a special combined Committee of the
Department of Agriculture and the pastoralists has been investigating the condition
for several years and experimental stations have been established in various parts
of the country, but no certain remedy has been discovered; the best results appear
to have been obtained by jetting and crutching under pressure, but this entails
constant care and treatment at intervals. Efforts to find.a deterrant which would
adhere to the sheep for any length of time have met with little success. Attempts
have been made to control the fly by means of liberating parasites, notably
Mormoniella brevicornis, but though success has been claimed in certain districts
this has not been the universal experience. More assistance might be obtained
by more careful attention to clearing up possible breeding places, particularly
of carcases, refuse around shearing yards, ete. In this connection the flight
experiments of Gurney become of importance as showing the extent of country
that may be infested by one dead sheep. Some six or more species ot fly have
been incriminated of attacking sheep, but some are probably merely secondary
invaders. The status of Ohrysomyia albiceps, for instance, as a sheep maggot fly
has been disputed, some authorities suggesting that it is predaceous on the larvae
of other flies present in the wool. It is probable, however, that this species does
actually attack the sheep, though it may also act as a predator.

The Oestridae or bot flies are represented in Australia by four introduced and
one native species. The habits of the introduced species appear to be identical
with their habits in Europe. Further information is, however, required in

PRESIDENTIAL ADDRESS. : XXV.

respect to the time of the year at which infestation occurs, as, according to
information gleaned by Mr. Clunies Ross in Western Australia, this appears to be
different in Western Australia from what it is in New South Wales. The one
native species, Tracheomyia macropi, is an inhabitant of the trachea of kangaroos;
only the larval and pupal stages are known; the adult does not appear to have
been seen.

Parasitic Flies: The Hippoboscidae are represented in Australia by three
introduced and a fairly large number of native species, most of which are as yet
undescribed. The introduced Hippobosca equina and H. camelina occur in
Western Australia, but are not known to carry any parasitic disease in Australia.
Both species are unrecorded from eastern States. The sheep ked, Melophagus
ovinus, is widely introduced. The native species mostly belong to the genus
Ornithomyia and are parasitic on birds; there are, however, three or four species
of Ortholfersia which are parasitic on marsupials (kangaroos and wallabies).

The Nycteribiidae or bat flies are represented by a number of species belonging
to the genera Nycteribia and Cyclopodia; the species have recently been mono-
graphed by A. Musgrave.

Streblidae are known to occur on Australian bats, but no species have so
far been described.

Siphonaptera: Australian fleas have been mainly investigated through the
researches of Dr. Karl Jordan and the late N. C. Rothschild. Some 35 species
are known, mainly parasitic on native rodents and the smaller marsupials. In
many instances the species are not restricted to one host, but may occur on a wide
range, including both rodents and marsupials. Bat fleas occur, but have not been
thoroughly studied. The Sarcopsyllidae are represented by some half-dozen species,
including the well known Echidnophaga gallinacea, the stick-fast flea which has
been the cause of considerable economic loss in poultry in Western Australia. It
is generally assumed that this species was introduced into Western Australia, but
it was certainly present in that State for several years before it attacked the
poultry and it is possible that the species is a native of Western Australia.
E. gallinacea was originally described from Ceylon, but it was already present as
a pest in that country and may well have been introduced from Western Australia.
Undoubtedly the home of the genus Echidnophaga is in Australia, to judge from
the number of species there represented. EH. gallinacea occurs also in South
Australia, but not commonly, whereas in Western Australia almost every animal
is affected and even children are attacked.

In addition to the native species of Siphonaptera, several introduced species
occur in Australia, of which the most important are the rat fleas, parasitic on the
introduced black and grey rats—R. rattus and R. norvegicus—and in particular the
so-called Indian rat flea—Xenopsylla cheopis—the vector of plague from rat to rat
and from rat to man. The association of this species with the transmission of
plague was first suggested by Simond in 1898, but it was Dr. Ashburton Thompson
of Sydney who, during the 1900 outbreak, clearly showed on epidemiological
grounds that the bubonic plague could be spread only through the medium of
such an insect as the rat flea. His conclusions were fully verified experimentally
by the Indian Plague Commission, of which Professor C. J. Martin, formerly of
Sydney and Melbourne, was a member.

The question of the distribution of Xenopsyllia cheopis in Australian ports is
of some importance. The species is known to occur abundantly on rats in Sydney,
Brisbane and other New South Wales and Queensland ports and in Perth, Western

XXvi. PRESIDENTIAL ADDRESS.

Australia. In Sydney about 45% of the rat fleas belong to this species and it
shows a distinct seasonal prevalence, being most abundant during the plague
epidemic months, February to May. The species occurs in Adelaide, but I have
no knowledge of its relative prevalence; whether it occurs in Melbourne, apart
from specimens taken on ship rats, appears uncertain. It seems to be absent
from Tasmania. It is noteworthy that Melbourne, Adelaide and Hobart have
always escaped infection at times when plague was rampant in other Australian
ports. The number of fleas captured from rodents in Sydney is shown on the
following table:

TABLE II.—FLEAS FROM RODENTS.
MICROBIOLOGICAL LABORATORY, 1909-1926.

Name 1909 1910) LOTT | 1912) 1913) 1974 TQ |) ALB. ||) TLS) L7 | 1918
2 eee pepe re ij
Xenopsylla | . |
OBOE 65 | oo) |l-ileat 701 | 1,105 641 213 233 22 300 197 | 67
Leptopsylla |
musculi 514 541 | 1,065 442 113 eal 156 108 60 | 63
Ceratophyllus |
fasciatus 210 266 375 PANG) 31 94 72 76 57 | 47
Ctenocephalus |
canis or felis.. 10 8 mal 14 3 2 4 8 1 1
Pulex irritans .. 1 aL 1 — — — — — — | —
Pygiopsylla
rainbowi — — = = as a= = — pees | =
Echidnophaga
myrmecobii = = ==> = = — cae = = | =
|
Total ILS PAD |) dS |) Gee) WSUS | 360 700 484 492 | 315 | 178
| | |
Name 1919 1920 | 1921 | 1922 | 1923 | 1924 1925 | 1926 Pee %
eS = |
Xenopsylia | |
cheopis 64 302 612 | 1,268 895 539 304 del || 9,215 || 44-55
Leptopsylla |
musculi 7 202 780 NG, 672 639 689 362 | 7,541 || 36°48
Ceratophyllus
fasciatus 30 169 503 542 376 284 309 107 || 3,767 || 18:22
Ctenocephalus
canis or felis.. 1 6 18 29 if 4 2 2 135 0-65
Pulez irritans .. == 1 2 4 2 4 — 16 0-08
Pygiopsylla |
rainbowi = — — — | =: 1 —— — 1 0-005
Echidnophaga | |
myrmecobii — — — 1 a — — 1 0-005
Z. pas | ee
Total 102 680 | 1,915 | 2,601 | 1,954 11,469 | 1,308 802 |/20,676 ||100-000
| | |

Hemiptera: The great order of the Hemiptera or bugs calls for little mention.
The common bed bug—Cimez lectularius—is introduced and is now widespread

throughout Australia.

It is not known to carry any disease in Australia, but its

PRESIDENTIAL ADDRESS. ; XXVIi.

presence is regarded as evidence of dirty and insanitary conditions. In Australia
we are fortunately free from such blood-sucking Hemiptera as Conorhinus and
its allies, which carry Chagas disease in South America.

Anoplura: The three species of human lice occur in Australia as well as the

common lice of domestic animals. The three great louse-borne infections of man—
Typhus, Relapsing Fever and Trench Fever—do not now exist in Australia, though
it is probable that typhus did occur in the early days of colonization. Reference
might be made to the occurrence in recent years of a typhus-like disease in
Adelaide. The disease is characterized by many of the features of typhus, though
in a milder degree, and the serum of patients gives the typical agglutination
reaction with Proteus X19. The disease is probably a member of a group of closely
allied diseases which includes typhus. The interesting point from the entomo-
logical aspect is that the disease does not appear to be infectious, but is probably
conveyed through the medium of some insect, or possibly acarine, parasite;
research has, however, failed so far to reveal the mode of transmission. A similar,
if not identical, disease has recently been recorded as occurring on farms in the
vicinity of Toowoomba, Q., and it is probable that cases elsewhere may have been
overlooked or confused with typhoid fever.
_. Other diseases exist in Australia, of which the aetiology and mode of
transmission is still unknown. Some of these may perhaps be insect- or mite-
borne. Mention has already been made of short-period coastal fevers in the
tropical zone; two other diseases as yet unelucidated might be mentioned—
Sarona Fever and Mossman River Fever. The latter at any rate appears to be
allied to Japanese River Fever and, like it, is probably carried by some species of
mite. :

The above review of medical and veterinary entomology in Australia does
not pretend to be exhaustive. Ample has, however, been said to show the
importance of the subject and something of the problems that still await
elucidation.

Hymenoptera: Though not containing any disease-carrying insects, the
Hymenoptera (bees, wasps and ants) are of interest to the medical and veterinary
entomologist from two points of view—firstly, many of the species will attack
human beings, inflicting severe stings; and secondly, several of the smaller
Hymenoptera are parasitic on flies of economic importance.

Stings from certain wasps, bees and ants are of common occurrence, but
are rarely recorded in medical literature. Cleland, in his paper “On Injuries
to Man due to Insects’, has collected a number of records. The more common
stinging ants belong mainly to the Ponerinae and include bull ants, soldier ants
and jumpers. A common stinging species is the greenhead (Chalcoponera
metallica). Several instances are on record where injured people have been
attacked by ants and portions of their faces or limbs eaten away.

The parasitic Hymenoptera of interest to us mostly belong to the Chalcidoidea.
Mormoniella brevicornis (Nasonia brevicornis) has already been mentioned; it is
parasitic in the pupae of blowflies and attempts, largely unsuccessful, have been
made to control the sheep-blowflies by its use. Other species have been suggested
for the same purpose, but have not been tested out on the same scale. Attempts
are now being made in Western Australia to establish the English blowfly
parasite (Alysia manducator). The Australian Hymenoptera have been fairly
extensively studied; the wasps by Dodd, Girault, Turner and others; the bees by
Cockerell, and the ants by Wheeler. Lately J. Clark, formerly of Western

XXVIii. PRESIDENTIAL ADDRESS.

Australia and now entomologist to the National Museum, Melbourne, has taken
up the study of the ants and has published several papers on the Ponerinae.

In conclusion, I desire to express my appreciation of the cordial co-operation
accorded me by my colleagues on the council and in particular by the permanent
officials—Dr. A. B. Walkom and Dr. G. A. Waterhouse. I also wish to thank
Mr. Carter and Dr. Mackerras for their help in revising the manuscript of this
address; the latter has been of great assistance in checking all references at a
time when I was unable to do so.

The following letter was sent to Their Royal Highnesses the Duke and Duchess
of York on the occasion of their visit to Sydney:

“We, on behalf of the Members of the Linnean Society of New South Wales—
a Society founded in 1874 to encourage the study of Natural History in Australia—
desire to offer to Your Royal Highnesses this brief record of our loyalty to the
Throne and of the pleasure evoked by the visit of Your Royal Highnesses as the
representatives of our revered Sovereign and Her Majesty the Queen.

May we be permitted to offer our respectful greetings and sincere wishes for
an enjoyable visit, and for the welfare of Your Royal Highnesses during your
return journey to Great Britain.”

It was with deep regret that we learnt of the death last’ week of Professor
A. A. Lawson, D.Sc., F.R.S., F.R.S.E. Professor Lawson was the first occupant
of the Chair of Botany in the University of Sydney, being appointed to that position
in 1913, when the Chair of Biology was divided, the late Professor W. A. Haswell
remaining as Professor of Zoology. The late Professor Lawson worked hard to
bring the School of Botany to its present high state of efficiency and had only
recently been elected a Fellow of the Royal Society of London. Professor Lawson
was a member of the Council of this Society in the years 1922-1924.

Dr. G. A. Waterhouse, Hon. Treasurer, presented the balance sheets for the
year ending 31st December, 1926, duly certified as correct by the Auditor, Mr. F.
H. Rayment, F.C.P.A., Incorporated Accountant; and he moved that they be
received and adopted, which was carried unanimously.

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

President: Professor L. Harrison, B.A., B.Sc.

Members of Council: C. Anderson, M.A., D.Sc., Professor A. N. St. G. H.
Burkitt, M.B., B.Sc., H. J. Carter, B.A., F.E.S., Sir T. W. E. David, K.B.E., C.M.G.,
D.S.0., B.A., D.Se., F.R.S., A. G. Hamilton and A. H. S. Lucas, M.A., B.Sc.

Auditor: F. H. Rayment, F.C.P.A.

The following resolution was carried unanimously: That the nerinere of the
Linnean Society of New South Wales assembled at the Fifty-second Annual General
Meeting desire to express their warm appreciation of the services rendered to the
Society by the retiring President (Dr. E. W. Ferguson) during his year of office
and their deep sympathy with him in his long illness and trust that he will make
a speedy recovery.

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JOSEPH JAMES FLETCHER.

Joseph James Fletcher was born in Auckland, New Zealand, in 1850. His
father, the Rey. J. Horner Fletcher, was then headmaster of what afterwards,
under his charge, developed into the ‘Auckland College’. He was both a
scholar and a theologian. Later on, for reasons of health, he resigned the head-
mastership and was appointed to the charge of a church at New Plymouth and,
after the outbreak of the Maori War in 1860, was transferred by the Methodist
Conference to Australia, first to Brisbane and then to Ipswich, where his son
gained a scholarship in the Grammar School. He was then appointed Principal
of Newington College Sydney, and there Fletcher passed most of his school days.
From school J. J. Fletcher went to the University of Sydney, where he took
his B.A. in 1870 and subsequently his M.A. degree in 1876. He does not seem to
have distinguished himself especially in the class lists, nor had he the physique
necessary for strenuous athletic work, but the following extract from The Sydney
Morning Herald of January 25, 1868, shows not only that he was a fellow student
of men who afterwards held prominent positions in the Commonwealth, but that
he must have had qualities that attracted the attention of that fine scholar and
experienced Professor, Dr. Badham. It reads as follows: “The Duke of Edinburgh
in New South Wales—Dramatic performance at the University. Mr. Cooper’s
(later Chief Justice of Queensland) impersonation of Phormio was very heartily
appreciated and so, too, was the Geta of Mr. (later Sir Hdmund) Barton and the
Nausistrata of Mr. Fletcher. This gentleman’s impersonation of the angry and
jealous wife elicited much merriment. Altogether Phormio was a decided success”.
Writing many years later Fletcher remembers how, “when the play was over and
we had been recalled and got our bouquets and as the curtain went down finally,
Dr. Badham came up out of.the prompter’s box and expressed his satisfaction at
the way things went off and gave me a word of commendation, having to speak
a foreign tongue in a big hall, in a falsetto voice’. He adds, “that made up to me
for all the grind of rehearsals. I did not get home to Parramatta till after
midnight, but I knew that my mother would sit up to hear whether I nad got a
bouquet and an encore, or had disappointed Dr. Badham’’. This little episode
occurred during his last year at the University. It is perhaps rather difficult to
recognize in the ardent naturalist of later years the Nausistrata of ‘“Phormio”.

After taking his degree he decided to devote himself to teaching, and
accepted a position in Melbourne on the staff of Wesley College, the Headmaster of
which was then Professor M. H. Irving, who had resigned the chair cf Classics
in the University to devote himself to school work. We know little of his work
in Melbourne except that, during the time he spent there, he came once more
under the influence of the distinguished head of a Public School. It was, however,
during this transitional period that he became interested in Natural Science, and,
in 1876, he resigned from Wesley College and left for London. This was the
time when Huxley was perhaps at his zenith—the year of his famous American
addresses. At South Kensington in Huxley’s lecture room and laboratory it
was possible for a student to listen to the leading exponent and protagonist of the

evolution theory, and at the same time gain by practical experience a first
c

Xexexive J. J. FLETCHER.

hand knowledge of plant and animal morphology. The latter is now a matter
of everyday experience in modern universities, but at that time anything like
true biological training was in its infancy. In the South Kensington laboratories
Fletcher met and worked with G. B. Howes, who later on succeeded to the
Professorship, and Jeffrey Parker, who migrated to New Zealand as Professor of
Biology in the Otago University. In those days, when the field of scientific
knowledge was comparatively limited, a Chair of Biology included teaching in both
Zoology and Botany, mainly from the morphological side. Whilst in England he ~
studied hard and systematically. There was then no such thing as recognition
of work done in Australian universities and, in order to gain the B.Sc. of the
London University, he had to start again, complete the full three years of work
and pass all the examinations necessary for admission to the degree. To start
again, however, is perhaps not strictly true because in Sydney, under Dr. Badaam,
his training, fortunately for himself, had been mainly on the literary and classical
side—a training that stood him in good stead in later years. He not only worked
in London but came into contact with F. Maitland Balfour, who, at Cambridge,
was laying the foundation in England of the science of Embryology. Struck with
his keenness and capacity Balfour invited him to visit the University as his guest
and there he spent three months amongst a band of students including Milnes
Marshall, Haddon, Lister, Adam Sedgwick, McBride, and others who in after
years occupied Chairs of Zoology and Biology in many British universities both
at home and in the Dominions. It was perhaps the most inspiring and fruitful
time in the history of Biological Science. Darwin, Huxley, Hooker and Wallace
were all at work, the “Challenger”, laden with wonderful material, had returned
after its three years’ cruise of investigation in southern and tropic seas and in
study, field and laboratory scores of eager students, relieved from the dead
weight of the special creation theory, were working under the stimulus of an
entirely new outlook on the world of life. It is difficult for students of the present
day to realize the excitement of those times when everything was new and stimu-
lating and when, further still, it was possible for one man to have a good all
round knowledge of, at all events, the salient features of different branches of
Science, small departments of which now occupy fully the time of many
investigators.

Imbued with the idea that to an Australian the study of problems concerned
with the Australian fauna and flora was a duty of paramount importance, he set
to work and, before leaving England in 1881, published in conjunction with J. J.
Lister who was then a demonstrator in comparative anatomy in the Cambridge
University, his first paper entitled “On the conditions of the median portion of
the vaginal apparatus in the Macropodidae’. The authors based their results
upon the scanty material then available in Cambridge and at the Zoological
Gardens, London, some of which had already been investigated by workers, such
as Owen and Everard Home, with indecisive results. In those days it was almost
impossible to devote one’s life to scientific work unless endowed with private
means, which Fletcher was not. Lack of pence, the attractions of his home
country, the hope to be able to do something towards the elucidation of its
fauna and flora whilst earning enough by other work to make this possible, drew
him back to Australia, where he spent the rest of his life.

Before leaving England, being fully aware of the imperfections of the
Australian libraries in regard to scientific literature, he set to work and compiled
a “Catalogue of Papers and Works relating to the Mammalian orders, Marsupialia,

J. J. FLETCHER. XXXV.

and Monotremata”’. The value of this, in those early days, especially to investi-
gators in Australia, was much enhanced by his inciusion of references to all the
new species described since the earlier works of Gould and Waterhouse. This was
published in Sydney in 1881. Immediately on arrival in Australia, personally and
with the aid of friends up country whom he interested in the work, he collected
much more material which served as the basis for further investigations into the
structure of the reproductive organs of Marsupials, and for the publication of three
additional papers in 1881 and 1888. His first three memoirs embody the results
of pioneer investigations into a matter of peculiar interest in regard to the
anatomy of Marsupials and are of permanent value. It is much to be regretted
that in those days there was no opening in Australia for a young unendowed
zoologist. No post was available, either for teaching or for carrying on systematic
research under modern laboratory conditions. Fletcher was obliged to return
to school work and thus Australia lost the services of one who would have been
a brilliant anatomist and an inspiring university teacher had the opportunity
presented itself.

He rejoined the staff of Newington College; was acting Headmaster for a
short time and then became one of the senior teachers. “As a teacher’’ says one
of his old pupils, Professor Scott Fletcher, now Professor of Philosophy in the
Queensland University, he “showed great originality and achieved quite remark-
able success. He had his own ideas on the subject of education, and evidently
believed that a boy at school should be guided in the development of his moral
character as well as taught to acquire knowledge. . . The roars of laughter
that at times rang out from Fletcher’s class-room were not only the envy of
adjoining “forms” but were ample evidence that the well known austerity of his
rule was not unrelieved by a genuinely human touch that stirred both laughter
and loyalty . . . his method of imparting knowledge was no less original
and effective’. This, be it remembered, was forty years ago, before the days of
Diplomas in Education, and of attempts to teach teachers how to teach. “He made
it a rule’, says his old student ‘‘never to tell his pupils anything which they could
find out for themselves. The spirit of research which he himself had learnt from
Huxley he passed on to his successive classes. No matter what the subject was,
he set his scholars to work gathering the raw material of fact which his own
master-hand was later to round off into systematic knowledge. There is perhaps
one word only in which may be summed up both his discipline and his instruction
—sinecerity. He hated all humbug and shams, but he loved all that is true or
beautiful or good in nature, in literature and in human character”.

From 1881-1885 Fletcher worked at Newington College, but during these four
years two things happened that determined the future course of his life. First,
he joined the Linnean Society of New South Wales, and there came into contact
with Sir William Macleay, its founder and benefactor, and secondly he married,
and in his wife found a companion who, with quiet, ceaseless and unselfish
sympathy, devoted herself to him and to his life’s work. Those of us who
have had the privilege of their private friendship, understand what this meant
to him and to us.

These were the early days of the Linnean Society. Macleay was evidently
impressed with the enthusiasm and capacity, and probably also, the ideals of
Fletcher in regard to Natural History work in Australia, and in 1885 offered him
the position of Director and Librarian of the Society. In reply to Fletcher’s
acceptance of the position, Sir William wrote to him a short but characteristic

XXXVi. J. J. FLETCHER.

note dated April 14, 1885. It runs as follows: ‘‘My dear Sir, I was much pleased
by your letter of yesterday, accepting my offer re ‘Linn. Soc.’. I shall now arrange |
that the first charge on the £10,000 I leave as an endowment to the Linnean, shall
be the salary of the Director, at present I presume my promise will be sufficient
security. I hope that the New Year will see our house completed. At all
events I should wish your duties to commence then. Yours, William Macleay”.

On January 1, 1886, Fletcher entered upon his duties, and henceforth devoted
his life to the work of the Society, to the furtherance of the study of Natural
History in Australia, and to the carrying out of what he conceived to be the aim
and objects of Sir William, and later also of Lady Macleay, with both of whom
he always remained on terms of the most friendly mutual trust and confidence,
which found expression in their selection of him as executor under both their wilis.

From 1886 to 1893 he acted as Director and Librarian; in 1893 the title of his
post was changed to that of Secretary in which capacity he acted until his retire-
ment in 1919. He was an ideal Secretary and, if the Society owes the possibility
of the present extent of its usefulness to the munificence and public spirit of
Macleay it owes almost an equal debt to the loyal and tireless zeal of Fletcher.
No task, however great or however small, that he thought touched upon the
honour of the Macleays or the welfare of the Society was left undone, no matter
what it cost him in time and thought. His width of scientific knowledge, his
literary ability, his wonderful acquaintance with work published in regarec to
Australian Natural History, and the early settlement of the Continent, together
with his long practical experience in the field, both as regards Zoology and
Botany, were of inestimable value to the Society and its members during the
thirty three years in which he held office and, it is not too much to say, largely
guided its destinies.

During his term of office he edited thirty-three volumes of proceedings with
meticulous care. Every word of every manuscript was carefully read through and
even references were verified before it went to the press, often indeed, in the case
of younger workers, before it was submitted to the Society, and many of them
gladly recognize that they owe him much for hints and suggestions derived from
his wide knowledge and experience.

Realizing, so far as he himself was concerned, the urgent need of studying
the land and fresh-water fauna before it passed away or became profoundly
modified by the opening up of the country and the introduction of alien forms, he
decided to work upon such material as he could study during leisure hours
without the necessity of recourse to a fully equipped laboratory. He started upon
the earthworms, a lowly group about which, in Australia, absolutely nothing was
known, though, in Europe, Darwin, in what is now regarded as a classic work, had
demonstrated their great practical value to the agriculturalist. From 1886-1894
he published six papers in which he showed the unexpected richness of Australia
in this group. He described some seventy new species belonging to nine genera,
five of which, as determined by him, were new and confined to Australia. His
work was not simply systematic, but included careful field observations in regard
to habits and distribution, and the results of anatomical investigation. “On one
occasion, when trying to estimate the number of worms in a given area”, he says,
“when the length of the furrows was 80 yards, I walked behind the plough and
counted all the worms I could see, either in the furrows or sticking out of the
overturned clods, and I found that for a number of furrows the average number

J. J. FLETCHER. XXXVii.

of worms under these circumstances was about 50 per furrow . . . these would
give nearly 10,000 worms to the acre’. At the same time he was working, together
with Mr. A. G. Hamilton, on the specific identity and distribution of Peripatus, and
was investigating the Land Planarians, studying them in their living conditions.
Of the latter only four species had previously been described, one by Darwin and
three by Moseley. Hight new species of Geoplana were added and six of
Rhynchodemus, a genus not before known from Australia.

He then turned his attention to Amphibia, more especially those to be
found near Sydney and in the County of Cumberland, where he was able to
watch them in their native state. His first paper dealt with their oviposition and
habits; the dates and places of spawning; the form of the spawn and the
dependence of this, in regard to its developing, upon the nature of the season. He
drew attention to the fact that certain species had developed the faculty, some in
the adult and others in the larval state, of accommodating themselves to the
varying climatic conditions of the dry parts of Australia. To those of us who
knew him and his methods of patient inquiry, the following sentences, quoted
from one of his papers, explain much. He says: “My most instructive round in
one of the suburbs of Sydney included a visit to an old quarry, a brickyard, a
deserted tanyard and three waterholes in paddocks used for watering cattie;
these five spots were frequented during some period of the year by at least eleven
species of frogs’. “That our frogs’, he says, “aestivate during hot and very dry
periods there can be no doubt. During such times one hears no croaking and
sees very little or nothing of the frogs, while logs and stones no longer afford
sufficient moist shelter. In March, 1885, a very dry month, after just sufficient
rain to moisten the ground, hearing croakings emanating from what, under more
favourable conditions, is the bed of a pond, I turned up the soil with a stick and
soon unearthed half a dozen specimens of Pseudophryne bibronii which were in

this manner trying to survive the drought. . . . P. australis is a lively perky
little frog, very partial to damp shelves and cracks in the Hawkesbury Sandstone
and breeds earlier. . . . P. bibronii on the other hand is much less active,

usually makes little or no effort to escape when uncovered in its hiding place,
‘shams dead’ when placed on its back and falls to the bottom like a stone when
thrown into water; I have never found it except on the ground, under stones,
logs, etc.; I have found the ova every year for seven successive years, once in
April only, thrice in May only, once in June only and twice in both April and
June”. These records and many others made by Fletcher are reminiscent of the
simple observations of a naturalist that, when recorded by White of Selbourne,
made him famous. Fletcher learnt to distinguish all the commoner frogs around
Sydney by their call. Sitting in the evening on his verandah at Hunter’s Hill,
enjoying with him the beauty of the fairy like scene of the Parramatta River,
framed with the delicate tracery of the trees in his garden, watching the ferry
boats gliding to and fro with long reflected lights of white and red and green, he
would listen for them, and when, from a pond close by, the shrill piping of the little
Hyla ewingi was heard, supported later by the slow-timed, dignified croak of its
larger relative Hyla cerulaea and the calls of two or three other species, he would
tell us which note belonged to each. At these times, while we could hear them
croaking in the water, he very much enjoyed repeating, by way of warning, a
sentence penned and published in an unlucky moment by a well known Professor
of Natural Science who had never worked in the field, “In Australia the Hylas
inhabit the tops of the lofty gum trees”.

XXXViii. J. J. FLETCHER.

He devoted in all seven papers to a careful investigation into the distribution
and habits of Australian frogs, and to the descriptions of new genera and species,
aided in this and other work by friends such as Messrs. Sidney Olliff, C. T. Musson,
W. W. Froggatt, A. M. Lea, A. G. Hamilton, T. G. Sloane and others who
recognized in him an ardent naturalist and were themselves as eager as he was
to elucidate the Australian fauna. His papers on the Batrachia are not only
models of accurate records of distribution but contain valuable original observa-
tions, such for example as those on the warning colouration of Notaden bennettii,
the Catholic frog.

The following, written privately but not published, shows his broad outlook
on problems concerned with the origin of the Australian fauna: ‘I feei satisfied
that the Tasmanian frog fauna is the most primitive assemblage of genera and
species (i.e. of frogs) we know of and that not only the genera but also the
species (with perhaps one or two exceptions) were differentiated in Antarctica.
Both in the case of Marsupials (extinct especially) and frogs and other things, I
cannot help thinking that there was much more differentiation in Antarctica than
we have been disposed to allow. Australia was a residuary legatee but the legacy
was of course subsequently increased by endemic differentiation. The chief
obstacle in the way seems to be the absence of intermediate forms of so many
groups. If the creatures were developed in the centre or on the Hast Coast (of
Australia) from forms which passed through Tasmania and Victoria, where are
they now? Or is it that we cannot recognize them when we see them?”

In January, 1900, he presided over the Biology Section at the Melbourne
meeting of the Australasian Association for the Advancement of Science, and
chose for the subject of his address “‘The Rise and early Progress of our knowledge
of the Australian Fauna”. He devoted to its preparation all his spare time during
the preceding eighteen months. “I can promise you’, he writes, “that there is
one President with his coat and weskit off and his pantaloons tied round with
string just below his knees—mentally perspiring—I hope to some purpose, if there
were not about 1,500 books that I want to see and don’t know how to find”. The
result was the compilation of a record of great value to all students interested
in the historic aspect of Natural History in Australia. “I have often’’, he says, ~
“wondered why no one seems to have attempted an historical sketch of the Fauna
such as I contemplated. Idiot that I was and am, I attempted to step into the
breach. Already I am wiser and sad and no longer wonder . . . the history
of the collections is most lamentably and grievously and sorrowfully depressing.

The collectors (in Australia) were full of enthusiasm, but the British
zoologists did not come up to the scratch. One is reminded of poor old W. B.
Clarke taking home his fossils—hawking them round the United Kingdom, going
down on his knees, begging and praying the British Palaeontologists to describe
them—getting nothing but kicks and cold shoulders. Determined to get them
worked up, he at last carts them off to Belgium, to De Koninck, who helps him
out of the difficulty”’.

In 1893 he edited the Macleay Memorial Volume, containing thirteen papers
on zoological, botanical and ethnological subjects, written by Australasian workers,
and himself contributed the biographical sketch. Writing in September of that
year he says: “We have got to the end of the lane at last, and the volume will
be published next week. I cannot say that I am proud of the sketch or that it does
justice to the theme. It cost an awful lot of trouble but the absence of journals,
letters and documents of that sort and the non-survival of hardly a contemporary

J. J. FLETCHER. XXXix.

with anything but shadowy recollections left me with but a small basis to work
on”’.

Times were then very straightened, financially, for the Linnean, as for all
other scientific societies in Australia, and he writes: “‘The situation at present may
be summed up in very few words—we hope soon to sell more copies and in the
meantime we have got an overdraft’. The latter caused him much anxiety. His
whole life was centred in the Society: indeed he was quixotic in a way that some-
times made it difficult for his colleagues on the Council to help him as they were
only too anxious and ready to do.

The following years were filled with routine work, and the endeavour to deal
with and solve, in a way satisfactory to the Society and in accordance with the
intentions of Sir William Macleay, serious legal and financial difficulties that arose
in connection with the foundation of the Research Fellowships and the appoint-
ment of a Bacteriologist. During all this time, however, he was working when-
ever opportunity offered, in close contact with friends and colleagues in the
Society—J. H. Maiden, A. H. S. Lucas, R. H. Cambage, H. J. Carter, C. T. Musson,
Henry Deane and others—each of whom was interested in some special branch put
recognized in Fletcher a naturalist of broad outlook and wide sympathies.

As time passed by he found it increasingly difficult, as a private worker with
no laboratory or assistance, to indulge in the rather expensive hobby of collecting
and maintaining a “spirit” collection that required constant supervision, and
therefore he devoted his energies more and more to botanical work, and to the
study of certain characteristic features of the Australian flora. Mr. C. T. Musson,
his friend and companion on many expeditions, in collaboration with whom he
worked for many years, says: “of late his time was chiefly devoted to Botanical
matters. He was especially keen about native things, the sandstone plants coming
first. He was always emphasizing the fact that this was a dry country and
that the plants had to be prepared, even down to the water’s edge for droughty
conditions. The amount of detail he got together when studying a subject was
remarkable. To give an example, when working at a Grevillea, he covered 50 or 60
quarto sheets with detail, hundreds of figures merely dealing with lobing of the
leaves. . . . His knowledge of the literature of our Fauna and Flora was
profound and his memory extraordinary”. During his work on Australian Acacia
phyllodes, the results of which were published in his Presidential Address in 1920,
he brought together, as every naturalist visitor to his house knew well, a vast
amount of material in specimens and notes. These led him eventually to the
conclusion that “The so-called phyllodes of the Australian phyllodineous Acacias
are not simply flattened petioles which have lost their blades . . . on the
contrary they are the flattened primary leaf axes or common petioles of bipinnate
leaves which have lost ‘their pinnae. . . . I propose to call them Euphyllodia
or Euphyllodes in the sense that they are something more than is implied in the
accepted definition of phyllodes”’.

Another important piece of work was carried out in conjunction with Mr.
Musson on shoot-bearing tumours of HEucalypts and Angophoras, published in
1918. Mr. Musson writes: “About eighteen years ago during a walk at the
Hawkesbury Agricultural College, we met with an extensive crop of seedlings of
Hucalyptus sideroxylon. On digging some up Fletcher remarked, on observing
the remarkable woody growths at the base of the stem, ‘A mallee in miniature’. In
their joint memoir the tumour was shown to originate in the axil of the cotyledons,
or in a few pairs of leaf axils immediately above them, from proliferating cambium

al J. J. FLETCHER.

material. The late Mr. J. H. Maiden first suggested a bacterial origin and in this
conclusion the authors concurred, the tumours, according to them, being due
to infection by some parasitic soil organism. In some species, spoken of as
‘refractory’, the growth slows down, a mild attack runs its course without injury
to growth. In the mallee gum the concrescence of the nodular growths is
complete; they enclose the tap and other roots and intercept water that they
-contain at the expense of the seedling stem; the so-called ‘mallee root’ is not a
root, but, in its extreme form, a great tumour, from which apparently, though in
reality only enclosed by it, stems and roots arise’.

A very characteristic sentence in one of his letters shows the way in which he
thought about such problems: ‘‘When the mallee scrubs have been cleared away
what are the soil organisms going to do? Are they going to send in their
resignations and die out, or are they going to adapt themselves to attacking the
introduced plants that have supplanted them?” After referring to Fletcher’s
work on Grevillea hybrids, for which he had fortunately prepared notes sufficient
to allow of its completion by Mr. Musson, the latter says: “He took his last walk
with me a fortnight before passing away. We sat in the park near the beautiful
Gordon Gully, and discussed various matters, chiefly dealing with Nature he loved
so well”.

The secretarial work of the Society, made even unnecessarily arduous by his
personal supervision of the minutest detail, began to tell upon him. Writing at
the close of 1918 he says: “I have not had a holiday since Easter, 1915, and my
eyes and brain are very tired, and I am retiring from the Linnean on 31st March,
1919, and have enough proof and indexing to keep me busy, so I enjoyed a week’s

loaf visiting my Grevillea hybrids. . . . I began to realize that thirty-three
years was going to be about as much as was good for me. So I decided to give
notice of my retirement. . . . I am normal again and looking forward to

finishing up properly, to leaving my work up to date and then having a holiday”.
He then refers appreciatively to the generous treatment accorded to him by the
Council; he himself, with characteristic modesty, had asked for a year’s retiring
allowance.

At the general meeting in March, 1919, the President, Professor H. G.
Chapman, expressed the deep appreciation of the Society of the distinguished
services that he had rendered to it. ‘‘No Society’, he said, “has received better
service than that given by our Secretary. The welfare of our Society has been
the sole care of his industry”. Dr. T. Storie Dixson, an old friend and past
President who had known Sir William Macleay in the early days, referred to the
Secretary’s loss as in many ways irreparable, especially by reason of the complete
understanding that existed between him and the founder in all matters concerning
its policy. Mr. A. H. S. Lucas, another past President, old friend and companion
in the field, handed over to the President a portrait to be hung in the Hall; Mr.
C. Hedley on behalf of the members presented a desk and chair to him, whilst
others bore testimony to the services rendered not only to the Society but to
themselves personally in connection with their work. To one of Fletcher’s
retiring disposition it was a rather trying ordeal but he felt deeply the genuine
expressions of appreciation and gratitude. He writes: “My comrades gave me a
very cordial and handsome send-off . . . half a dozen speeches and a resolution
passed. The warmth and enthusiasm of the proceedings fairly astonished me as I
had no knowledge beforehand of this part of the ceremony (referring to the desk

J. J. FLETCHER. xli.

and chair) and they pretty nearly knocked me off my perch, but I managed to hang
on and say what it was needful for me to say without being overcome’’.

He was elected President for the years 1920 and 1921. In his first address
he dealt with the two main forms of climatic conditions with which men on the
land are concerned. He pointed out that the man in the northern hemisphere has
learnt his lesson and can live in harmony with his environment. The winter
conditions when he must house and feed his stock are so regular that he can
arrange his programme by the almanac. The man in Australia has yet to learn
how to adapt himself to Nature’s second method for giving the land its needed
rest and sweetening by means of what he aptly called a periodic “drought sleep’.
We have manuals of Faunas and Floras, of fodder grasses and minerals and so on,
but how is it, he asked, that we have no manual of drought problems to teach man
how, with the aid of knowledge gained from scientific investigation, to insure
against damage by drought. In this address he included as a supplement the
results of his research, spread over many years of field work, into the so-called
Phyllodes of Acacias.

At a special general meeting held in June, 1920, to commemorate the centenary
of the birth of Sir William Macleay, he delivered an address on “The Society’s
Heritage from the Macleays’. In regard to this he wrote: “I will show my
Macleayan things that I am using as the basis for my address for William Macleay’s
centenary on June 13th next. Some original drawings of Lewin, W.S.M.’s (William
Sharp Macleay) original sketches, one of ‘an animal caught by Mr. Huxley in the
tow net in Torres Straits’, portraits, letters, etc. Lady Macleay gave me these when
she went to England about 25 years ago. I looked over them, saw they were
interesting, iocked them up, wondering how I could use them. Now I know what
they are and what they mean, and I am up to my neck in it. When I was proposed
as President I did not want it. Then I thought of the centenary and how I could
use the material for an address—‘The Society’s Heritage from the Macleays’.” The
address, which occupied 68 pages of the Proceedings, contains a most valuable
record of the lives and work of the three senior members of the family, Alexander,
William Sharp, and Sir George Macleay; of the collections they brought together;
of their scientific contemporaries; of their 97 years’ connection with the parent
society in London; of their 51 years’ connection with the Australian Museum. It
might more correctly be entitled: ‘Australia’s Heritage from the Macleays”’. It
was only his inside knowledge of the family, gained as the result of the complete
trust placed in him by Sir William and Lady Macleay, who gave him access to
their private journals and papers, that made it possible for him to compile such a
valuable historical record of early days not only of the Society but in tke Colony.
The most pleasing feature to him was the presence of Admiral Dumaresq, of
H.M.A.S. “Melbourne”, then in Sydney, a grandson on his mother’s side of
Alexander Macleay. The Admiral was deeply interested in the relics shown, and
at his invitation Fletcher spent some interesting hours with him on his Flagship,
showing him letters and papers and discussing the early days of the Macleays,
within sight of the ancestral home garden at Elizabeth Bay.

In his Presidential Address for 1921 he referred to a subject in regard to which
he felt very keenly, that a grave injustice was being done to the memory and
generosity of the Macleays, and to the public interested in Natural History. Under
the heading of “Is all well with the Macleay Museum at the University?’ he
certainly showed that it was far from well and that the University was not either
in letter or in spirit carrying out its share of the bargain. It was the only time

D

xlii. J. J. FLETCHER.

Fletcher entered into a newspaper controversy, but he felt that “one of Sir
William’s great enterprises, potentially so fructifying, if properly managed has
become bankrupt”, and he was quite right. When it was over he wrote, “very busy
and very tired and in need of a holiday. At all events what I have done I have
done from a sense of duty”.

During his thirty years at Elizabeth Bay he had accumulated a very valuable
collection consisting especially of marsupial material, amphibia, earthworms, and
Peripatus. It included all the material upon which his zoological work was based.
In 1923 he spent months in overhauling, rearranging and labelling it and finally
presented everything to the Australian Museum. It pained him not to be able
to present it to the Macleay Museum, but he was not prepared to place it, as he
wrote rather scathingly, “in a room, placed under an iron roof, no blinds to the
windows, the curator not a zoologist”.

After retirement from active work at Elizabeth Bay he spent his spare time
in carrying on botanical work on Acacias, Grevilleas, and Loranthaceae, often again
in company with Linnean friends of long standing—Messrs. J. H. Maiden, R. H.
Cambage, G. H. Halligan and C. T. Musson. Especially was he fond of exploring
botanically the Hawkesbury sandstone and rarely missed a weekly tramp along
the upper reaches of Lane Cove, the beauty of which, as seen in later years, from
his own verandah, was a never failing source of pleasure by day and night. He
writes in 1922, when he was a free man, “Of late the sunsets, sunrises, golden
smokes, golden mists and reflections have been particularly beautiful. It is true
that more and more houses are going up, and more and more bush is going, but
I return thanks daily that though A may own this bit of ground and house, and B
that bit, yet none of them, nor all of them own the Lane Cove, or the reflections,
or the landscape”.

He naturally felt the removal of the Linnean offices to the new premises in
College Street, and whilst it was in progress wrote: ‘‘Before the Society can
vacate Elizabeth Bay and remove the Library, a new one will have to be built.
Then, if possible, the Bay property will be offered for sale. You can imagine
what a wrench all this is and promises to be to me. It will certainly be more
convenient for members and for meetings and is perhaps to be regarded as
inevitable, but it is a little sad for the small remnant of the old brigade to cut
ourselves adrift from the memories and traditions of the bit of the old historic
garden and our associations with Sir William, and the old surroundings”.

As the result of an accident four years ago he was forced to abstain from
anything like strenuous exertion, though still retaining his association with the
Society, and keen interest in his own work and that of his friends, who looked
forward to his being able to enjoy some years of well earned rest in surroundings
that appealed to him, both because of their scientific interest and of their beauty.
But it was not to be, and on Saturday, May 15, 1926, without any warning, he
passed away suddenly. It may be truly said that he lived for the Society and
the carrying on of the Macleay tradition and, as the resolution of the Council
records, “happily lived to see its present high position with an already honoured
tradition behind it and an ever increasing activity in scientific research of the
highest standard well established and portending a prominent future, with which
his name will ever be inseparably associated’’.

W.B.S.

ABSTRACT OF PROCEEDINGS.

SPECIAL GENERAL MEETING.
30th Marcu, 1927.

Mr. A. G. Hamilton, Vice-President, in the Chair.

Pursuant to notice the Honorary Treasurer moved the following addition to
Rule vi: The Honorary Treasurer is authorized to accept from any ordinary
member, who is not in arrear, the sum of Fifteen Guineas in lieu of further annual
subscriptions.

The motion was seconded by Professor Harrison and carried unanimously.

ORDINARY MONTHLY MEETING.
30th Marcu, 1927.

Professor L. Harrison, B.A., B.Sc., President, in the Chair.

The Donations and Exchanges received since the previous Monthly Meeting
(24th November, 1926) amounting to 61 Vols., 603 Parts or Nos., 24 Bulletins, 12
Reports and 14 Pamphlets, received from 160 Societies and Institutions and 7
private donors were laid upon the table.

PAPERS READ.

1. Notes on Australian Diptera. No. x. By J. R. Malloch. (Communicated
by Dr. E. W. Ferguson.)

2. The Anatomy of Cheilanthes vellea. By May M. Williams, M.Sc.,
Linnean Macleay Fellow of the Society in Botany.

3. A New Deltopecten from the Illawarra District, N.S.W. By John Mitchell.

4. The Fossil Estheriae of Australia. Part i. By John Mitchell.

5. The Geology of the Country between Lamb’s Valley and the Paterson
River. By G. D. Osborne, B.Sc.

6. Note on a Dicotyledonous Fossil Wood from Ulladulla, N.S.W. By C.
Barnard, B.Sc.

7. On a Case of Natural Hybridism in the Genus Grevillea (Proteaceae).
By C. T. Musson and the late J. J. Fletcher, M.A., B.Sc.

NOTES AND EXHIBITS.
Miss H. Claire Weekes, B.Sc., Linnean Macleay Fellow of the Society in
Zoology, contributed a preliminary note on placentation in some lizards. Definite
placentation amongst lizards has been described in Chalcides tridactylus and
C. ocellatus by Giacomini (1891) and (1906) respectively; in Tiliqua scincoides
by Flynn (1923); and in Lygosoma (Liolepisma) entrecasteauxzi by Harrison and
Weekes (1925).
The occurrence of omphaloplacentation and allantoplacentation in the Scincid
lizards Lygosoma (Hinulia) quoyi, Egernia whitei and E. striolata and of
B

xliv. ABSTRACT OF PROCEEDINGS.

omphaloplacentation in TJ. scincoides is here recorded for the first time. In
Hinulia quoyi the allantoplacentation is highly specialized. In the classic
C. tridactylus, T. scincoides and L. entrecasteauxi the foetus obtains its food from
the parent by means of the glandular activity of the modified uterine epithelium
and the absorbing and phagocytic powers of the enlarged chorionic ectoderm cells.
In other words it is the epithelial tissues which play the important part in food
transition. In H. quoyi the epithelial tissues partly degenerate and there is a
concentration of maternal and foetal capillaries in close apposition. Hence it can
be stated that the type of allantoplacentation found in H. quoyi more closely
resembles that found in the Marsupialia than does any hitherto recorded in a
Reptile. In H#. whitei and LE. striolata the allantoplacentation resembles that of
H. quoyi but offers sufficient variation to be of interest.
Detailed accounts of these phenomena are in course of preparation.

SPECIAL GENERAL MEETING.
27th Apri, 1927.

Professor L. Harrison, B.A., B.Sc., President, in the Chair.

The minutes of the Special General Meeting of 30th March, 1927, were read
and confirmed.

The President pointed out that the addition of Rule vi as carried and confirmed
at the Special General Meetings provides for Life Membership of the Society.

ORDINARY MONTHLY MEETING.
27th Aprin, 1927.

Professor L. Harrison, B.A., B.Sc., President, in the Chair.

Mr. W. A. L. Bredero, Orange, Mr. T. G. Campbell, Manly, Mr. W. Dixson,
Killara, Miss M. L. Garde, Neutral Bay, and Mr. K. C. Richardson, B.Sc., Sydney,
were elected Ordinary Members of the Society.

The President announced that. Messrs. A. F. Basset Hull, R. H. Cambage,
C.B.E., F.L.S., H. J. Carter, B.A., F.E.S., and Dr. E. W. Ferguson had been elected
Vice-Presidents; and Dr. G. A. Waterhouse, Hon. Treasurer for the Session 1927-28.

The President drew attention to the Fourth International Congress of
Entomology to be held at Ithaca, New York, in August, 1928.

The Donations and Exchanges received since the previous Monthly Meeting
(30th March, 1927) amounting to 7 Vols., 71 Parts or Nos., 7 Bulletins, 2 Reports
and 3 Pamphlets, received from 52 Societies and Institutions and 3 private donors
were laid upon the table.

PAPERS READ.

1. Two New Species of Setaria from Western Australia. By A. S. Hitchcock.
(Communicated by Mr. W. M. Carne.)

2. The Influence of certain Colloids upon Fermentation. Parts iv, v and vi.
By R. Greig-Smith, D.Sc., Macleay Bacteriologist to the Society.

3. Notes on Australian Mosquitoes (Diptera, Culicidae). Part i. The
Anophelini of the Mainland. By I. M. Mackerras, M.B., Ch.M., B.Sc., Linnean
Macleay Fellow of the Society in Zoology.

4. The Physiographic and Climatic Factors controlling the Flooding of the
Hawkesbury River at Windsor. By Lesley D. Hall, B.Sc.

ABSTRACT OF PROCEEDINGS. xlv.

NOTES AND EXHIBITS.

Mr. C. T. Musson exhibited specimens of the common introduced snail Helix
aspersa from Gordon, Sydney, and also for comparison specimens from England.
He remarked that he could see little difference in their texture.

Mr. John Mitchell exhibited a very small cylindro-conical stone from an
aborigines’ camp and workshop on the seashore south of the Bellambi jetty. In
general contour it resembles the stones of this character optained from the
western parts of New South Wales, but differs from all of these in its small size
(four inches long) and from most of them in being smooth and having a rounded
base (Mem. Geol. Surv. N.S.W., Ethnol. Ser. No. 2). The stone in question, it is
plain, has been made from a thin pebble of some igneous rock, ground into its
present shape. It is sub-two-sided; one side is flat, the other convex; each end is
rounded. He was not aware of any previous reference having been made to the
discovery of a similar stone fashioned by the aborigines of the east coast of
Australia.

Mr. J. Mitchell also exhibited a fragment of fossil fish belonging to the genus
Elonichthys, recently obtained by him from the insect beds near Belmont, N.S.W.
Very few fossil fish have been found in the Newcastle Coal Measures. The first
was Urosthenes australis Dana; the next was not found till 1912, when he
secured a few specimens, and now the present specimen. The latter specimens

belong to the genus Elonichthys, which seems to be the genus best represented
in the Newcastle Coal Measures.

ORDINARY MONTHLY MEETING.
25th May, 1927.

Professor L. Harrison, B.A., B.Se., President, in the Chair.

Mr. J. W. F. Armstrong, Bogan River, N.S.W., and Mr. C. G. Oke, St. Kilda,
Victoria, were elected Ordinary Members of the Society.

The President announced that a movement had been started to provide a
suitable memorial to the late Professor A. A. Lawson, the first Professor of Botany
in the University of Sydney.

The President announced that the Society had that day vacated the Hall at
Elizabeth Bay which it had occupied since 2nd January, 1886. The Library had
been removed to the Macleay Museum, University of Sydney, where it would
remain until it could be housed in its new home.

The Donations and Exchanges received since the previous Monthly Meeting
(27th April, 1927) amounting to 25 Vols., 370 Parts or Nos., 31 Bulletins, 7 Reports
and 19 Pamphlets, received from 73 Societies and Institutions and 2 private donors
were laid upon the table.

PAPERS READ.

1. The Interpretation of the Radial Field of the Wing in the Nematocerous
Diptera, with Special Reference to the Tipulidae. By C. P. Alexander.
(Communicated by Dr. E. W. Ferguson.)

2. New Gall-forming Thysanoptera of Australia. By Dudley Moulton.
(Communicated by Mr. W. W. Froggatt, F.L.S.)

38. Note on Reproductive Phenomena in some Lizards. By Miss H. Claire
Weekes, B.Sc., Linnean Macleay Fellow of the Society in Zoology.

xIvi. ABSTRACT OF PROCEEDINGS.

4. An Ecological Study of the Flora of Mt. Wilson. Part iv. Habitat ;
Factors and Plant Response. By J. McLuckie, M.A., D.Sc., and A. H. K. Petrie,
M.Sc.

NOTES AND EXHIBITS.

Mr. David G. Stead referred to an extraordinary occurrence of large sharks
in the following note: Several species of large sharks are known to occur in
Port Jackson. Some of these penetrate but a short distance from the sea and
only one kind travels far up into the head waters. On the 24th instant a launch
party under the charge of Mr. Charles Messenger was fishing along the eastern
channel of the harbour entrance when they met with a large school of sharks.
Hight of these were captured and were seen by me to-day. They were all of the
species Carcharinus brachyurus Gunther, the Whaler or Mullet Shark, and ranged
in size from about 7 feet 6 inches to 10 feet. Apparently all of those present in
this school were of the same species. This appears to be the first record of the
occurrence of the Whaler in school formation. Both the Grey Nurse, Carcharias
arenarius Ogilby (with which the present specimens were confused in the daily
press accounts) and the well-known Blue Pointer are known to occur in very
large schools on “outside” grounds, but the Whaler is usually more diffused—
existing singly or in pairs ({ and 2). Mr. Messenger, who is a highly experienced
shark catcher, is of the opinion that had several boats been present not iess
than one hundred sharks might have been taken. The Whaler is the most
dangerous of our man-eating sharks, penetrating our estuaries, even up into
fresh water. It commonly attains to 10 feet in length and is of great girth.
The occurrence of this large school of Whaler sharks is probably an aftermath of
the schooling season for Sea Mullet, Mugil cephalus Linnaeus.

Dr. C. Anderson communicated the following note:—At a meeting of this
Society held on 27th June, 1888 (These ProceEepines, 1888, p. 894), Dr. J. C. Cox
exhibited “a Tertiary fossil from Wildhorse Plains, which he believed to be
identical with Thylacodes decussatus Gm., a living Port Jackson species”; this
name is synonymous with Vermetus decussatus, a marine mollusc with a vermiform
shell. By some unfortunate mistake this note led to the tentative recognition of a
new genus of fossil marsupial, ? Thylacodes Cox, which is listed in Roger’s
“Verzeichniss der bisher bekannten fossilen Saugetiere” (Ber. d. naturw. Vern. f.
Schwaben u. Neuburg, Augsburg, 1896) in the family Phalangistidae, and in
Trouessart’s “Catalogus mammalium tam viventium quam fossilium’’ (Vol. II,
1898-9, p. 1156) in the family Phalangeridae, sub-family Thylacoleontinae, along
with Thylacoleo carnifex Owen, the so-called Marsupial Lion. In the Supplement
to this latter work (1904-5, p. 846) this curious mistake is repeated but the
sub-family is transferred to the family Dasyuridae.

ORDINARY MONTHLY MEETING.
29th Jung, 1927.

Mr. R. H. Cambage, C.B.E., F.L.S., Vice-President, in the Chair.

The Chairman offered the congratulations of the Society to Dr. W. G.
Woolnough on his appointment as Acting Federal Geologist.

The Chairman announced with pleasure that the Government had recently
issued a Proclamation protecting certain of the wild flowers.

The Donations and Exchanges received since the previous Monthly Meeting
(25th May, 1927) amounting to 21 Vols., 106 Parts or Nos., 7 Bulletins, 7 Reports

ABSTRACT OF PROCEEDINGS. xlvii.

and 2 Pamphlets, received from 69 Societies and Institutions and 2 private donors
were laid upon the table.

PAPERS READ.

1. The Gasteromycetes of Australasia. vii. The Genera Disciseda and
Abstoma. By G. H. Cunningham.

2. The Gasteromycetes of Australasia. viii. The Genus Mycenastrum. By
G. H. Cunningham.

3. Additional Flora of the Comboyne Plateau, 1926. By E. C. Chisholm,
M.B., Ch.M.

4. Further Notes on a new Classification of Australian Robberflies (Diptera,
Asilidae). By G. H. Hardy.

5. The Phylogeny of some Diptera Brachycera. By G. H. Hardy.

NOTES AND EXHIBITS.

Mr. A. J. Nicholson exhibited from the Macleay Museum Collection a bilateral
gynandromorph of the butterfly, Papilio androgeus from S. America.

Mr. A. Musgrave exhibited, from the Australian Museum Collection, a remark-
able aberration of the female of Papilio aegeus from New South Wales. In the
specimen the red spots on the hindwing were much enlarged and drawn out te
about four times their usual size.

Dr. G. A. Waterhouse exhibited the following butterflies: (1) Mosaic
gynandromorphs of (a) Troides priamus pronomus from Cape York in which the
female markings predominated, (bv) Papilio aegeus ormenus from Darnley Is.
showing about equal proportions of male and female markings; (2) specimens of
Papilio oberon from Santa Cruz Is. recently caught by Messrs. E. Le G. Troughton
and A. A. Livingstone, and suggested that as they resembled Papilio aegeus from
Eastern Australia so closely, they may have been introduced into the Santa
Cruz Group.

Dr. Waterhouse also exhibited fossils from Brookvale, near Manly, consisting
of Estheria coghlani and a beetle. The latter, the first record of a beetle from the
Quarry, was recently found by Mr. W. H. Hatcher.

Mr. David G. Stead exhibited two extraordinary fishing lines: (1) A crocodile
line as used by the Malays of the Malay Peninsula, Southern Siam, Sumatra and
Borneo, for the capture of the short-snouted crocodile (Crocodilus porosus). The
line itself is formed of a thin, strong rotan (Mal.) cane, to which the unusual
hook is fastened—by the middle of the shank—by means of a multi-strand snood
of ramie fibre. The strands of the snood are quite separate so as to prevent the
crocodile from biting through the line, which becomes entangled in the teeth.
After the crocodile swallows the baited hook, the tension of the line on the
middle of the hook causes it to lie right athwart the gullet. (2) A baitless
and barbless hooked fishing line, known as rawai (Mal.) used for the capture of
sharks and rays and great eels by Chinese (Hokkien) fishermen. The line is set
chiefly at the edge of tidal flats on muddy and sandy bottoms.

Mr. E. Cheel exhibited specimens of Hremophila longifolia, infested with the
aecidial stage of a rust-fungus (Uredineae) collected at Thackaringa, Broken Hill,
by Mr. A. Morris. The cylindrical orange-red coloured pseudoperidia with white
tips somewhat agree with those described as Roestelia polita by Berkeley which
according to McAlpine (Rusts of Australia, p. 98) is an Uromyces. The only record
so far as he could ascertain of a Myoporaceous plant being found infested with

xviii. ABSTRACT OF PROCEEDINGS.

a “rust-fungus” is that of Cunningham (Trans. N.Z. Inst., 1924, p. 35) on
Myoporum laetum.

Mr. A. F. Basset Hull exhibited 103 examples of 40 species, 23 genera of
abnormal Loricates, showing specimens having 5, 6, 7 and 9 valves; others with
damaged and repaired valves, or with one side repaired, and the aperture on the
other side closed by enlargement of the adjoining vaive. The greater number of
these had been obtained since his exhibit before the Society in May, 1925.

Dr. I. M. Mackerras exhibited a series of Diptera Brachycera, showing South
American affinities.

Mr. R. H. Cambage recorded a series of flights of the flying squirrel Petauroides
volans, which in six successive glides covered a distance of 590 yards. A resident
at Milton, in the ’seventies, while near his house in the twilight, saw a squirrel
leave the top of a Hucalyptus tree 100 feet high and glide to the foot of another
70 yards away. This it immediately climbed and from its summit glided to the
next at 80 yards. It lost no time in ascending three more trees at distances of
110, 120 and 90 yards apart respectively, and from the top of the last it glided to
another 120 yards away, which it climbed and in which it remained, this
evidently having been its objective. While climbing the trees it uttered its
peculiar squealing call notes as if to give a friendly warning of its approach
to any of its kindred that might be in the tree. It is possible the squirrel landed
on the ground a few yards short of some of the trees, but its total journey of a
third of a mile shows that these nocturnal marsupials may wander a considerable
distance from their homes to which they return by the morning.

Archdeacon F. E. Haviland exhibited a device for use in centering ana
mounting microscopic objects. It consists of a wooden slab recessed, into whicn
fits a brass slab with a central hole and recessed for cover glasses. It is claimed
that the object is held in its intended position on the slide and air bubbles are
avoided.

Mr. G. P. Whitley exhibited specimens of the Surf Fish, Iso rhothophilus,
collected by Messrs. F. A. McNeill and A. A. Livingstone at Long Bay baths
on 5th May, 1927.

These fishes, which live in the outer breakers of the surf, occurred in
thousands in the baths, where they had evidently been stranded after the Easter
cyclone. Numerous isopod parasites were clinging to the fishes or swimming
freely amongst them, whilst some copepods were netted with the fishes, and had
evidently become detached from them. These crustaceans have been identified by
Mr. H. M. Hale as: One young female Irona, five stages of juvenile Cymothoid
isopods, and numerous Caligus sp. The collectors noticed some of the fishes
in a school turning sideways and sinking to the bottom, where many were
eaten by rock-pool fishes.

Subsequent observations made at intervals by Mr. Whitley showed that the
turning of apparently sickly fishes was not usually performed by those carrying
parasites, and that taken as a whole the fishes fell on their left or right sides,
but single specimens turned to right or left consistently. The reason for this
action is still unknown. The fishes were plentiful until the end of May, but on
5th June, practically all had disappeared. The majority probably returned to the
ocean in the waves which had broken over the baths during the stormy weather
meanwhile.

X-ray photographs of three specimens, made by Surgeon Lieut.-Commander
W.E. J. Paradice, R.A.N., were also shown. The vertebrae numbered 13 + 29 to 30

ABSTRACT OF PROCEEDINGS. xlix.

(excluding hypural). Iso flos-maris, known in Japan as the “Flower of the Wave,”
has 18+ 25 (Jordan and Starks).

Dr. McLuckie exhibited specimens of Chert obtained from the Rhynie Chert
bed, Scotland, discovered in 1913 by Dr. Mackie, containing the petrified remains
of very simple early Devonian plants. Fragments of the plant remains, sections
and photographs of sections were also exhibited. The fragments had been
separated from the main Chert mass by heating and suddenly cooling in cold
water. He gave a brief account of the plant remains petrified in this Chert,
discussing the extremely simple vascular structure and organization of Rhynia and
Hornea which were leafless and rootless. Absorption from the soil was carried
on by the rhizoids which develop from the creeping rhizome. The aerial branches
forked, bore stomata (and probably chlorophylliferous tissue) and terminal
sporangia, and had a very simple organized central vascular cylinder, consisting
of spiral or annular tracheids.

In Rhynia and Hornea we have the most simply organized vascular plants
known. The sporangial structure of Hornea with its central sterile tissue and
characteristic dome-shaped spore-bearing zone suggests a moss-like character (¢.g.
Sphagnum) and a certain analogy is apparent between the Mosses and these
simple vascular plants, which show a combination of characters which have been
stressed by different botanists to show their Pteridophytic, Thallophytic or
Bryophytic affinity. The Rhyniaceae may be a synthetic group related to Pterido-
phytes and Bryophytes and retaining some characters of their original Algal
ancestry.

The discovery of the Rhyniaceae and the excellent elucidation of their
structure by Lang and Kidston during recent years has given us a knowledge
of the remarkably simple and probably the most primitive of vascular plants. ‘They
throw a certain amount of light upon the probable origin of the sporophytic
generation of land plants.

ORDINARY MONTHLY MEETING.
27th JuLy, 1927.

Mr. H. J. Carter, B.A., F.E.S., Vice-President, in the Chair.

The Chairman referred to the great loss the Society had sustained by the
death of Dr. EH. W. Ferguson on 18th July and read the following resolution passed
by the Council and sent to Mrs. Ferguson on behalf of the Society: “The Society
places on record its very deep regret at the death of Dr. E. W. Ferguson. It
realizes the debt it owes to the late Dr. Ferguson, who had been a member of the
Society since 1908, a member of the Council since 1921, its President during 1926.
His death is a loss to the members of the Society and to the naturalists of
Australia by whom he was held in the highest esteem and respect. The Society
expresses to Mrs. Ferguson and her family their very sincere sympathy with them
in their sad bereavement’.

Mr. M. F. Albert, Elizabeth Bay; Miss M. V. McHugh, Potts Point; Rev.
H. M. R. Rupp, Paterson, N.S.W.; and Mr. W. L. Waterhouse, M.C., B.Sc.Agr.,
Roseville were elected Ordinary Members of the Society.

The Chairman offered the congratulations of the Society to Mr. A. H. K.
Petrie, M.Sc., on his appointment to an 1851 Exhibition Travelling Scholarship.

The Donations and Exchanges received since the previous Monthly Meeting
(29th June, 1927) amounting to 3 Vols., 105 Parts or Nos., 3 Bulletins and 20
Reports, received from 53 Societies and Institutions and 2 private donors were
laid upon the table.

1. ABSTRACT OF PROCEEDINGS.

PAPERS READ.

1. Australian Coleoptera: Notes and New Species. No. v. By H. J. Carter,
B.A., F.E.S.

2. The Gasteromycetes of Australasia. ix. Keys to the Genera and Species
of the Lycoperdaceae. By G. H. Cunningham.

3. Notes on Australian Diptera. No. xi. By J. R. Malloch. (Communicated
by Dr. I. M. Mackerras.)

4. Descriptions of New Species of Australian Coleoptera. Part xix. By
A. M. Lea, F.E.S.

5. The Vegetation of the Kosciusko Plateau. Part i. The Plant Communities.
By J. MeLuckie, M.A., D.Sc., and A. H. K. Petrie, M.Sc.

NOTES AND EXHIBITS.

Dr. G. A. Waterhouse exhibited bred specimens of (1) Ogyris oroetes from
the following new localities: Clermont, Queensland (EK. J. Dumigan), Brisbane,
Queensland (L. Franzen), and Belltrees, near Scone, N.S.W. (G.A.W.); (2) the
three subspecies of Ogyris amaryllis from all the States of Australia except
Tasmania; (3) specimens of Ogyris aenone taken near Cairns by Mr. A. N. Burns.

Mr. W. L. Wearne exhibited a piece of cedar timber in which a portion of a
bed-post was embedded. The annual rings showed that this had been in position
for about fifty years.

Mr. A. F. Basset Hull drew attention to the recent issue of “A Monograph of
the Australian Loricates’” and presented a copy to the Society on behalf of the
Royal Zoological Society of New South Wales.

ORDINARY MONTHLY MEETING.
3lst AuGUST, 1927.

Mr. R. H. Cambage, C.B.E., F.L.S., Vice-President, in the Chair.

The Chairman referred to the death on 6th August of Dr. R. Greig Smith,
Macleay Bacteriologist to the Society and read the following resolution passed by
the Council: “That the Council places on record its very deep regret at the death
of Dr. R. Greig Smith, who was the first Macleay Bacteriologist to the Society,
occupying that position for nearly thirty years. During the whole of his residence
in New South Wales he had devoted himself assiduously to his research work and
had assisted in many ways the advancement of science in Australia. His death is
a loss to members of the Society and his presence will be missed from its meetings.
The Society expresses to Mrs. Greig Smith its very sincere sympathy in her sad
bereavement”.

The Chairman also referred to the death of Mr. Thomas Whitelegge, at one
time a member of the Society and for several years a member of Council.

Mr. W. D. Francis, Brisbane, Queensland, was elected an Ordinary Member
of the Society.

The Chairman announced that Mr. A. G. Hamilton had been elected a Vice-
President and Mr. A. J. Nicholson, M.Sc., a member of Council.

The Chairman announced that the Société de Physique et d’Histoire Naturelle
de Genéve offers a prize for the best unpublished monograph on a genus or family
of plants, the closing date being 31st March, 1930.

The Donations and Exchanges received since the previous Monthly Meeting
(27th July, 1927) amounting to 31 Vols., 191 Parts or Nos., 20 Bulletins, 3 Reports
and 8 Pamphlets, received from 98 Societies and Institutions and 2 private donors
were laid upon the table.

ABSTRACT OF PROCEEDINGS. li.

PAPERS READ.

1. Notes on Australian Diptera. No. xii. By J. R. Malloch. (Communicated
by Dr. I. M. Mackerras.)

2. Notes on Australian Mosquitoes (Diptera, Culicidae). Part ii. The
Zoogeography of the Subgenus Ochlerotatus with Notes on the Species. By I. M.
Mackerras, M.B., Ch.M., B.Sc., Linnean Macleay Fellow of the Society in Zoology.

3. The Xerophytic Structure of the Leaf in the Australian Proteaceae.
Part i. By A. G. Hamilton.

4. Australian Hesperiidae. Part i. Notes and Descriptions of New Forms.
By G. A. Waterhouse, D.Sc., B.E., F.E.S.

NOTES AND EXHIBITS.

Dr. I. M. Mackerras exhibited a series of Australian Tachinidae in illustration
of Mr. Malloch’s paper.

Mr. A. G. Hamilton showed a number of lantern slides to illustrate the
xerophytic structure of the leaf of the Australian Proteaceae.

Dr. G. A. Waterhouse exhibited the new subspecies described in his paper and
a number of coloured lantern slides of Australian Butterflies.

Mr. David G. Stead exhibited a photograph showing bees collecting bitumen
from the coating of wood pipes at Yarrangobilly (N.S.W.).

Mr. Stead invited the attention of the Society to the great destruction of
Koalas (Native Bears) which had taken place in Queensland as a result of the
short open season of one month (August) declared by the Queensland Government.
Already many thousands of skins had actually been exported and thousands more
were in store in Sydney and Brisbane. On his motion it was resolved that the
Commonwealth Government be requested to refuse all applications for permits
to export such skins in the future in accordance with their proclamation of
5th December, 1923.

Mr. G. P. Whitley exhibited a sample of marine growth from off Long Reef,
New South Wales. This growth had been reported from Broken Bay, the ocean
off Long Reef, and local estuarine waters, where it had occurred as a kind of
scum at varying depths. It had proved annoying to fishermen by clinging to their
lines, becoming concentrated into lumps as the lines were pulled out of the water.
It has been suggested that the presence of this organism might have a deleterious
effect on marine animals and that its occurrence in such large quantities might
have been due to the lengthy dry weather period just experienced. It was
suggested that the organism is a marine alga.

Mr. A. H. S. Lucas exhibited specimens of the English primrose and cowslip
and also what was probably a hybrid between them. He pointed out that the
hybrid was found growing in a habitat between the two parent forms.

ORDINARY MONTHLY MEETING.
28th SEPTEMBER, 1927.

Mr. A. F. Basset Hull, Vice-President, in the Chair.

The Chairman referred to the death of Mr. Daniel Frederick Cooksey on 16th
September, 1927. He had been a member of the Society since 1926.

The Chairman drew attention to the cable from England announcing the death
of Professor A. Liversidge, who had been one of the original members of the
Society and had served on its Council during 1875 and 1876.

The Chairman announced that the Council is prepared to receive applications
for four Linnean Macleay Fellowships tenable for one year from ist March, 1928,

lii. ABSTRACT OF PROCEEDINGS.

from qualified candidates. Applications should be lodged with the Acting
Secretary, who will afford all necessary information to intending candidates, not
later than Wednesday, 2nd November, 1927.

The Donations and Hxchanges received since the previous Monthly Meeting
(31st August, 1927) amounting to 9 Vols., 87 Parts or Nos., 2 Bulletins and 3
Pamphlets, received from 44 Societies and Institutions and 3 private donors were
laid upon the table.

PAPERS READ.

1. Studies in the Goodeniaceae. Part i. The Life History of Dampiera stricta
R.Br. By P. Brough, M.A., B.Sc., B.Sc.Agr.

2. Notes on Australian Diptera. No. xiii. By J. R. Malloch. (Communicated
by Dr. I. M. Mackerras.)

3. Notes on Australian Marine Algae. Part iv. The Australian Species of the
Genus Spongoclonium. By A. H. 8. Lucas, M.A., B.Sc.

NOTES AND EXHIBITS.

Dr. I. M. Mackerras announced that the late Dr. E. W. Ferguson had
bequeathed his fine collection of Amycterids (Coleoptera, Curculionidae) to the
Macleay Museum, University of Sydney and that the collection had been transferred
to the Museum.

ORDINARY MONTHLY MERTING.
26th OctToBErR, 1927.

Mr. H. J. Carter, B.A., F.H.S., Vice-President, in the Chair.

Rev. E. Norman McKie, Guyra, N.S.W., was elected an Ordinary Member of the
Society.

Candidates for Linnean Macleay Fellowships, 1928-29, were reminded that
Wednesday, 2nd November, 1927, is the last day for receiving applications.

Mr. D. G. Stead mentioned the movement that had been proceeding for some
time for the appointment of a Royal Commission of Inquiry by the Commonwealth
Government, to make a close investigation into the general status of the aborigines
throughout Australia. Upon his motion, seconded by Mr. I. V. Newman, it was
resolved to write to the Prime Minister in support of the proposal.

The Donations and Exchanges received since the previous Monthly Meeting
(28th September, 1927) amounting to 12 Vols., 95 Parts or Nos., 13 Bulletins and
2 Reports, received from 56 Societies and Institutions and 1 private donor were
laid upon the table.

PAPERS READ.
1. Notes on Australian and Exotic Sarcophagid Flies. By G. H. Hardy.
2. Placentation and other phenomena in the Scincid Lizard, Lygosoma
(Hinulia) quoyi. By H. Claire Weekes, B.Sc., Linnean Macleay Fellow of
the Society in Zoology.

NOTES AND EXHIBITS.

Mr. D. G. Stead drew attention to the feeding habits of the large Black Shag,
Phalacrocorar carbo, in the lower waters of Sydney Harbour. After many years’
observation it was concluded that normally this species lived almost exclusively
upon bottom-frequenting fishes, especially the Frog Fish, Batrachus dubius, which

ABSTRACT OF PROCEEDINGS. liii.

was often obtained from a depth of about 26 feet. Catfish, Cnidoglanis megastoma,
and Flathead, Platycephalus spp., were also captured.

Mr. Stead also drew attention to the presence (in waters adjacent to Sydney
Heads) of the Wafer Worm, Leptoplana australis, a species of dendrocoelous
turbellarian worm which was destructive to oysters.

Mr. EH. Cheel exhibited specimens of two species of plants of the Selagineae
group of the family Scrophulariniaceae as follows:—(1) Hebenstreitia dentata
Linn. This is quite common in the neighbourhood of Newcastle and has evidently
been brought from South Africa in ballast. (2) Selago corymbosa Linn.
Hawkesbury Agricultural College, C. T. Musson. Probably introduced from South
Africa in agricultural seeds. It is interesting to note that Oenothera biennis
(Evening Primrose) is also quite common at Newcastle associated with Oenothera
odorata and Hebenstreitia.

Mr. T. H. Pincombe exhibited Cleithrolepis granulatus found (July, 1927) in
sewer tunnelling at the head of Tambourine Bay, Sydney, in a similar deposit to
that at Brookvale.

Dr. G. A. Waterhouse showed a lantern slide of the pupa of Liphyra brassolis
and gave some notes on its life history.

ORDINARY MONTHLY MEETING.
380th NoveEMBER, 1927.

Professor L. Harrison, B.A., B.Se., President, in the Chair.
The President extended a welcome to Dr. A. B. Walkom on his return from
' England.

Mr. Bert BertRAM, Greenwich, Mr. F. A. Crart, Ashbury, Professor T. G. B.
Ossorn, Adelaide, and Miss Doris A. SELBy, Gordon, were elected Ordinary Members
of the Society.

The President announced that the Council had reappointed Miss H. Claire
Weekes and Miss Ida A. Brown to Linnean Macleay Fellowships in Zoology and
Geology respectively for one year from 1st March, 1928.

The President offered congratulations to Dr. R. J. Tillyard on his appointment
as Chief of the Biological Research Station at Canberra; to Mr. C. A. Sussmilch on
his appointment as Principal of the East Sydney Technical School and Assistant
Superintendent of Technical Education; also to Mr. C. Barnard, B.Sc., on his
appointment as Assistant Botanist to the Council of Scientific and Industrial
Research.

The President announced the receipt of a formal acknowledgment from the
Secretary to the Prime Minister of the resolution carried at the last meeting.

The Donations and Exchanges received since the previous Monthly Meeting
(26th October, 1927) amounting to 216 Vols., 298 Parts or Nos., 14 Bulletins and
6 Reports, received from 75 Societies and Institutions and 1 private donor were laid
upon the table.

PAPERS READ.

1. Notes on Australian Marine Algae. No. vy. By A. H. S. Lucas, M.A., B.Sc.

2. Mosquito Control in the Municipality of Lane Cove, New South Wales.
By B. Bertram. (Communicated by Dr. I. M. Mackerras.)

3. A new Dendrobium for New South Wales and Queensland. By Rev.
H. M. R. Rupp.

liv. ABSTRACT OF PROCEEDINGS.

NOTES AND EXHIBITS.

The President (Professor L. Harrison) exhibited examples of a new species
of the primitive Annelid Stratiodrilus from the gill cavities of the Madagascar
crayfish Astacoides madagascariensis. Through the courtesy and kindness of the
Consul-General for France, M. Nettement, and the Governor-General of Madagascar,
specimens of this crayfish were forwarded to Miss Lucy M. Wood, B.A., who
desired to examine them for Temnocephaloid parasites, and who was successful in
obtaining these.

The genus Stratiodrilus was established by the late Professor W. A. Haswell,
F.R.S., in 1900, for a species obtained from the Tasmanian crayfish Astacopsis
tasmanicus. In 1913 he described a second species from the Australian crayfish
Astacopsis serratus. In 1921 a third species was discovered in Uruguay by Dr.
E. H. Cordero, which has not so far been described. The present species makes the
fourth.

The discovery is of considerable interest, as it sheds some light on the origin
of the northern and southern crayfishes, a subject which has formed the basis for
classical discussions at the hands of Huxley, Ortmann, Géoffrey Smith, Matthew
and others.

It is now established that both Temnocephala and Stratiodrilus occur in South
America, Australia and Madagascar, chiefly as ectoparasites of crayfishes, while
neither occurs in the northern hemisphere. Yemnocephala has no close relations,
but the only known relative of Stratiodrilus, Histriobdella homari occurs upon
the Norway lobster, Nephrops norvegicus. The suggestion is tentatively made that
both northern and southern crayfishes have been derived from Nephrops-like
marine ancestors, but that the transition from marine to freshwater conditions
was. in the northern hemisphere, too sudden for Histriobdellid parasites to survive
it; while in the southern hemisphere a more gradual transition has allowed
survival. Temnocephala would appear to have developed entirely as a freshwater
parasite, possibly derived from a Rhabdocoele ancestry.

Mr. A. Musgrave exhibited a coloured drawing of an Argiopid spider,
Poecilopachys bispinosa (Keyserling), made by Miss Joyce Allan of the Australian
Museum, from a specimen collected by Mr. B. Bertram at Lane Cove, Sydney.
During life the colour of the spider was observed to undergo changes similar to
those in the octopus, the colour ebbing and flowing along the fore-border of the
abdomen and the abdominal spines, and producing a definite flush. The abdominal
spines, too, underwent a change, being quite smooth at times, while at other times
they would be papillose. The spider is not uncommon in the vicinity of Sydney.
It has been recorded from Gayndah, Queensland, and the Island of Upolu, Samoa.

DONATIONS AND EXCHANGES. ly.

DONATIONS AND EXCHANGES.

Received during the period 25th November, 1926, to 30th November, 1927.
(From the respective Societies, etc., unless otherwise mentioned.)

ABERYSTWYTH.
Welsh Plant Breeding Station, University College of Wales.—Advisory Bulletin

No. 2 (1927); Bulletin, Series H, Nos. 5-7 (1927); “Self and Cross-
fertilisation in Lolium perenne L.”, by T. J. Jenkin (From Journ. of Genetics,
xvii, 1, Aug., 1926); “The Welsh Journal of Agriculture’, iii (1927).

ACCRA.
Geological Survey of the Gold Coast.—Report for the Period April, 1925-March,

1926 (no date).

ADELAIDE.
Department of Mines: Geological Survey of South Australia—Annual Report

of the Director of Mines and Government Geologist for 1926 (1927); Mining
Review for the Half-Years ended June 30th, 1926 (No. 44) (1926);
December 31st, 1926 (No. 45); June 30th, 1927 (No. 46) (1927).

Field Naturalists’ Section of the Royal Society of South Australia.—‘The
South Australian Naturalist’, vii, 4 (1926); viii, 2-4 (1927). :
Public Library, Museum and Art Gallery of South Australia.—Forty-third

Annual Report of the Board of Governors for 1926-1927 (1927); “List of
Books on Ornithology in the Public Library of South Australia and other
Adelaide Libraries” (1926); Records of the South Australian Museum, iii,

38, (1927).

Royal Geographical Society of Australasia, South Australian Branch.—
Proceedings, xxvii, Session 1925-26 (1927).

Royal Society of South Australia.—Transactions and Proceedings, 1 (1926).

South <Australian Ornithological Association—“The South Australian
Ornithologist’, ix, 1-4 (1927).

University of Adelaide——‘The Australian Journal of Experimental Biology and
Medical Science’, iii, 4 (T.p.&c.) (1926); iv, 1-3 (1927); “The Pink Boll-
worm of Queensland”, by F. G. Holdaway (From Bull. Ent. Res., xvii, 1,
1926); Commemoration Address, 1925—“Some Aspects of Forestry in South
Australia’, by Prof. Sir Douglas Mawson (1925); 27 Reprints from Trans.
Proc. Roy. Soc. 8S. Aust., xlix-]1 (1925-1926); Med. Journ. Aust., 1925 (1925);
Pap. Proc. Roy. Soc. Tas., 1925 (1925); and Rept. Aust. Asscn. Adv. Sci.,
xvii, 1924 (1926).

Woods and Forests Department.—Annual Progress Report for the Year 1925-26
(1926).

ALBANY.
New York State Library, University of the State of New York.—New York State
Museum Bulletin, Nos. 269-273 (1926-1927). 5

lvi. DONATIONS AND EXCHANGES.

ALGER.
Institut Pasteur d Algerie.—Archives, iv, 1-4 (T.p.&c.) (1926).
Société ad Histoire Naturelle de lVAfrique du Nord.—Bulletin, i, 1909/10—xvi,
1925, 1-7, 9 (1910-1925); xvii, 1926, 7-9 (Index) (1926-1927); xviii, 1927,
1-6 (1927).

AMSTERDAM.
Nederlandsche Entomologische Vereeniging.—Entomologische Berichten, vii,
150-155 (1926-1927); Tijdschrift voor Entomologie, Ixix, 3-4, Supplement
(T.p. & ec.) (1926); Ixx, 2 (1927).

ANN ARBOR.

University of Michigan.—Contributions from the Museum of Geology, ii, 7-8
(1926); Miscellaneous Publications of the Museum of Zoology, Nos. 15-17
(1926); Occasional Papers of the Museum of Zoology, Nos. 166-183 (1926-
1927); Papers of the Michigan Academy of Science, Arts and Letters, vi-vii,
1926 (1927).

AUCKLAND.
Auckland Institute and Museum.—Annual Report, 1926-27 (1927).

BALTIMORE.
Johns Hopkins University.—University Circulars, N.S., 1926, 2-11 (1926); List
of Publications, 1927 (1927).

BANDOENG.
Geological Survey in the Dutch East Indies.—Wetenschappelijke Mededeelingen,
Nos. 5-6 (1927).

BARCELONA.
Real Academia de Ciencias y Artes de Barcelona.—Memorias, xix, 15-17
(T.p.&c.) (1926); xx, 1-4 (1926-1927); Nomina del Personal Academico,
1926-1927 (1927).

BASEL.
Naturforschende Gesellschaft—Verhandlungen, xxxvii-xxxviii, 1925/1926-
1926/1927 (1926-1927).

BERGEN.
Bergens Museum.—Aarbok, 1926, 1-2 (T.p.&c.) (1927); Aarsberetning, 1925-
1926 (1926).

BERKELEY.

University of California.—Publications: Botany, xi, 3-7; xiii, 11-19; xiv, 1-2
(1926-1927) ; Entomology, iv, 2-5 (1926-1927); Geology, T.p. &c. for xv (1924-
1926); xvi, 5-12 (1926-1927) ; xvii, 1 (1927); Physiology, T.p. &c. for v (1915-
1926); vii, 1-2 (1926-1927); Zoology, T.p.&c. for xxi (1918-1926); and xxiv
(1922-1926); xxix, 8-18 (T.p.&c.) (1926-1927); xxx, 6-8 (1926-1927); xxxi,
1-5 (1927).

DONATIONS AND EXCHANGES. lvii.

BERLIN.

Botanischer Garten und Museum.—Notizblatt, ix, 89-90 (T.p.&c.) (1926-1927) ;
x, Oil CMAs

Deutsche Entomologische Gesellschaft, E. V.—Deutsche Entomologische
Zeitschrift, 1926, 3-5, Beiheft (T.p. & c.) (1926-1927); 1927, 1-2 (1927).

Deutsche Entomologische Museum.—Entomologische Mitteilungen, xv, 5-6
(T.p.&c.) (1926); xvi, 1-5 (1927); Supplementa Entomologica, Nos. 14-15
(1926-1927).

Notgemeinschaft der Deutschen Wissenschaft.—“Flora”’, Neue Folge, xxi, 2-4
(T.p. &c.) (1926-1927) ; xxii, 1-2 (1927).

BERN.
Naturforschende Gesellschaft.—Mitteilungen ad. Jahre 1926 (1927);
Verhandlungen, 1926 (1926).

BIRMINGHAM.
Birmingham Natural History and Philosophical Society.—List of Members,
1927, and Annual Report, 1926; Proceedings, xv, 6, Session 1926-27 (1927).

BomBay.
Bombay Natural History Society—GJournal, xxxi, 3-4 (1926-1927); T.p.&c. for
xxxi, pts. 1 and 2; xxxii, 1 (1927). -

Haffkine Institute—Reports for the Years 1925, 1926 (1926, 1927).

BONN.
Naturhistorische Verein der Preussische Rheinlande und Westfalens.—
Sitzungsberichte, 1925; Verhandlungen, 83, 1926 (1926).

Boston.
American Academy of Arts and Sciences.—Proceedings, lxi, 7-12 (T.p. &c.)
(1926); Ixii, 1-4 (1927).
Boston Society of Natural History.—Memoirs, viii, 2 (1916); Proceedings,
xxxv, 2-3 (1915); xxxvili, 4-7 (1926-1927).

BRISBANE.

Department of Agriculture and Stock.—Queensland Agricultural Journal, xxvi,
5-6 (T.p. &c.) (1926); xxvii, 1-6 (T.p.&c.) (1927); xxviii, 1-5 (1927).

Queensland Government Mining Journal (from the Editor).—xxvii, Dec., 1926
(T.p.&c.) (1926); xxviii, Jan.-Nov., 1927 (1927).

Queensland Museum.—Memoirs, ix, 1 (1927).

Queensland Naturalists’ Club and Nature-Lovers’ League.—Queensland
Naturalist, vi, 1-3 (1926-1927).

Royal Geographical Society of Australasia, Queensland Branch.—Queensland
Geographical Journal, N.S., xl-xli, 1924-26 (1926).

Royal Society of Queensland.—Proceedings, xxxviii, 1926 (1927).

BROOKLYN.

Botanical Society of America.—American Journal of Botany, xiii, 8-10
(T.p. & ce.) (1926); xiv, 1-8 (1927).

lviii. DONATIONS AND EXCHANGES.

BRUSSELS.

Académie Royale de Belgique.—Annuaire, 93rd Année, 1927 (1927); Bulletin
de la Classe des Sciences, 1926, 3-12 (T.p.&c.) (1926-1927); 1927, 1-3
(1927).

Société Entomologique de Belgique.—Bulletin and Annales, Ixy, 8 (1925);
Ixvi, 8-12 (Index) (1926); Ixvii, 1-8 (1927).

Société Royale de Botanique de Belgique.—Bulletin, lviii (1925); lix, 1-2
(T.p. &c.) (1926-1927).

Société Royale Zoologique de Belgique—Annales, lvi-lvii, 1925-1926 (1926-
1927).

BUDAPEST.
Musée National Hongrois.—Annales, viii, 1 (1910); xiii, 1 (1915); xxii-xxiv
(1925-1926).

BUENOS AIRES.
Museo Nacional de Historia Natural.—Anales, xxiv (19138); xxxiii (1923-1925) ;
Comunicaciones, ii (1923-1925); Memoria Anual de 1924 (1925).
Sociedad Argentina de Ciencias Naturales.—Revista “Physis’”, vii, 27 (T.p. &c.)
(1925); viii, 28-30 (1925-1926).
CAEN.
Société Linnéenne de Normandie.—Bulletin, 7° Série, viii, 1925 (1926);
Mémoires, xxvi, 2 (T.p.&c.) (1924); Nouvelle Série, Section Botanique, i,
1 (1926); Section Zoologique, i, 1 (1925).

CALCUTTA.

Geological Survey of India.—Memoirs, Palaeontologia Indica, N.S. iii, 4 (1911) ;
iv, 1-4 (T.p.&c.) (1911-1912); vi, 3 (1917); vii, 3 (1927); ix, 2 (1927);
x, 1-2 (1927); xiv, 1 (1927); Records, lix, 3-4 (T.p.&c.) (1926-1927); 1x,
1 (1927).

Indian Museum.—Records, xxviii, 3-4, Appendix and T.p.&c. (1926-1927) ;
xxix, 1-2 (1927); Report on the Zoological Survey of India for the Years
1923-1926 (1926).

CAMBRIDGE, England.

Cambridge Philosophical Society.—Biological Reviews and _ Biological

Proceedings, ii, 1-3 (1926-1927); Transactions, xxiii, 10-11 (1927).
CAMBRIDGE, Mass.

Museum of Comparative Zoology at Harvard College—Annual Report of the
Director for 1925-1926 (1926); Bulletin, Ixvii, 12-16 (T.p.&c.) (1926-1927) ;
Ixviii, 1-8 (1927).

CAPE Town.

Royal Society of South Africa.—Transactions, xiii, 4 (T.p.&c.) (1926); xiv,
1-3 (1926-1927); xv (complete) (1926).

South African Museum.—Annals, xxiii, 3 (T.p.&c.) (1926); xxv, 1 (1927);
Guide Leaflet No. 2 (1926); Guide to the Relief-Map of the South-western
Portion of the Cape Province (1926); Report for the Year ended 31st
December, 1926 (1927); Short Guide to the Museum (1926).

DONATIONS AND EXCHANGES. lix.

CHICAGO.

Chicago Academy of Sciences.—‘An Annotated Flora of the Chicago Region”,
by H. S. Pepoon (Bulletin viii, Natural History Survey) (1927).

Field Museum of Natural History.—Department of Anthropology, Guide, Part 6,
Ethnology of Polynesia and Micronesia by Ralph Linton (1926); Publica-
tions, Botanical Series, iv, 5 (1927); vi, 1 (1926); Report Series, T.p. and
Index to Illustrations for vi, 1921-1925; vii, 1 (1927); Zoological Series,
xii, 12-13 (1926); xiii, 5 (1927); xvi, 1-2 (1926).

CHRISTCHURCH.
Philosophical Institute of Canterbury.—Transactions and Proceedings of the
New Zealand Institute, lvi-lvii (1926-1927).

CINCINNATI.
Lloyd Library.—Bulletin No. 27 (1926); Mycological Notes, Nos. 73-75 (1924-
1925).

CLugs, Roumania.
Gradina Botanica.—Bulletin, vi, 1-4, Appendix 1, T.p. &c. (1926) ; vii, Appendix 1
(1927); Contributions Botaniques de Cluj, i, 8 (1927).

Comp Spring HaRpor.
Station for Experimental Evolution, Carnegie Institution of Washington.—
Annual Report of the Department of Genetics, 1925-1926 (Extracted from
Year Book No. 25 for the Year 1926) (1926).

CoIMBRA.
Universidade de Coimbra: Instituto Botanico.—Boletim da Sociedade Broteriana,
Serie 2, iv (1927).

CoLOMBO.
Colombo Museum.—Spolia Zeylanica (Ceylon Journal of Science, Section B—
Zoology and Geology), T.p. &c. for xiii (1924-1926); xiv, 1 (1927).

CoLtumBus, Ohio.

American Chemical Society.—Industrial and Engineering Chemistry, xviii,
10-12 (T.p.&c.) (1926); xix, 1-8, 10 (1927); News Edition, iv, 20-24 (Index)
(1926); v, 1-20 (1927).

Ohio Academy of Science and Ohio State University Scientific Society.—
Ohio Journal of Science, xxvi, 5-6 (T.p. &c.) (1926); xxvii, 1-4 (1927).

Ohio State University Ohio State University Bulletin, xxxi, 5 (1926).

COPENHAGEN. -
Det. Kgl. Danske Videnskabernes Selskab.—Biologiske Meddelelser, v, 8-9
(T.p. &c.) (1926); vi, 1-7 (T.p. &c.) (1926-1927); Oversigt, Juni, 1925-Maj,
1926 (1926); Juni, 1926-Maj, 1927 (1927).
Zoological Museum of the University.—Publications, Nos. 44-48 (1926); The
Danish Ingolf-Expedition, iv, 6 (1926); v, 10-11 (1926-1927).

DUBLIN.
Royal Irish. Academy.—Proceedings, xxxvii, Section B, Nos. 10-13; 16-21 (1926-
1927).
Cc

lx. DONATIONS AND EXCHANGES.

DURBAN.
Durban Museum.—Museum Report for the Municipal Year ended 31st July,

1926 (1927).

East LANSING.
Michigan Agricultural College, Agricultural Experiment Station.—“A Rapid

Method for Performing the Agglutination Test, Etc.”, by I. F. Huddleson
and E. R. Carlson (Reprinted from Journ. Amer. Vet. Med. Asscn., 1xx, N.S.
23, No. 2, November, 1926); “Further Studies on the Isolation and Cultiva-
tion of Bacterium Abortus (Bang)’’, by I. F. Huddleson, D. E. Hasley and
J. P. Torrey (Reprinted from The Journal of Infectious Diseases, xl, 2,
February, 1927); Report of the Veterinary Division for the Year ending
June 30, 1925 (no date).

EDINBURGH.
Royal Physical Society.—Proceedings, xxi, 2, Session 1925-26 (1926).
Royal Society of Edinburgh.—Proceedings, xlvi, 3-4 (T.p.&c.), Session 1925-
26 (1926-1927); xlvii, 1-2, Session 1926-27 (1927); Transactions, liv, 3
(T.p. & c.), Session 1925-26 (1926); lv, 1, Session 1926-27 (1927).

FRANKFURT ON MAIN.
Senckenbergische Naturforschende Gesellschaft—Abhandlungen, xl, 2 (1927);
Berichte, 56, 10-12 (T.p. &c.) (1926); 57, 1-6 (1927).

FREIBURG, I. Br.
Naturforschende Gesellschaft.—Berichte, xxvi, 2 (T.p.&c.) (1926); xxvii, 2
(Mp &e:) (1927).

GENEVA.
Société de Physique et d'Histoire Naturelle.—Compte Rendu des Séances, xliii,
33 (Abo, C2@,)) (EPA Ss oxbhie al (alka ye

GENOA.
Museo Civico di Storia Naturale.—Annali, 1, 1921-1926 (1921-1926).
Societa Entomologica Italiana.—Bollettino, Anno lviii, 8-10 (T.p.&c.) (1926);
lix, 1-4, Supplement (Memoria, v, 1, 1926), 5-7 (1927).

GIESSEN.
Botanische Institut.-—‘Uber Vitalfarbung von Pflanzenzellen mit Phthaleinen’’,
by E. Kuster (Bericht Ober. Gesell. fur Natur- und Heilkunde, Neue Folge,
Bd. xi) (1926).

GRANVILLE.
Denison University—Journal of the Scientific Laboratories, xxi, pp. 285-434
(T.p. &c.) (1926); xxii, pp. 1-135 (1927).

HAMBURG.
Naturwissenschaftlicher Verein.—Verhandlungen, Vierte Folge, ii, 2-4
(T.p. &e.), 1925 (1925).

DONATIONS AND EXCHANGES. lxi.

HELSINGFORS.

Societas pro Fauna et Flora Fennica.—Acta, liv (1924-1926); Ivii, 1 (1927);
Acta Zoologica Fennica, 1-2 (1926-1927); Memoranda, 1-2, 1924/1925-
1925/1926 (1927).

Societas Scientiarum Fennica.—Acta, 1, 7-15 (T.p.&c.) (1926); Arsbok-
Vuosikirja, iv, 1925-1926 (1926); Bidrag, 80, 6-7 (1926-1927); Commenta-
tiones Physico-Mathematicae, ii, 19-30 (T.p.&c.) (1926); iii (1926); Tables
Générales des Publications, ii, 1911-1926 (1927).

Societas Zoolog.-botanica Fennica Vanamo.—Annales, iii-v (1926-1927).

HoBaRt..

Department of Mines, Geological Survey.—Geological Survey Record, No. 4
(1915); Geological Survey Report Nos. 4, 5, 7 (1914, 1915); Report of the
Director of Mines for Year ending December 31, 1926 (1927); Report of the
Secretary for Mines for the Years ending December 31, 1914-1925 (1915-
1926).

Royal Society of Tasmania.—Papers and Proceedings for the Year 1926 (1927).

Tasmanian Field Naturalists’ Club.—The Tasmanian Naturalist, N.S. ii, 2
(1927).

Hono.uuu, T. H.
Bernice Pauahi Bishop Museum.—Bulletins 29-40 (1926-1927); List of Publi-
cations, No. 5 (August, 1927).
Pan-Pacific Union— Bulletin, N.S. Nos. 83-86, 88, 89 (1926-1927); “Journal of
the Pan-Pacific Research Institution’, ii, 1-4 (1927).

INDIANAPOLIS.
Indiana Academy of Science.—Proceedings, 35, 1925 (1926).

IrHaca, N.Y.
Cornell University.—15 Separates (1919, 1926-1927); “Landbouonderwys deur
die Skool”, by F. E. Geldenhuys (Bloemfontein, 1925).

JAMAICA PLAIN.
Arnold Arboretum.—Journal, vii, 3-4 (T.p.&c.) (1926); viii, 1-3 (1927).

JOHANNESBURG.
South African Association for the Advancement of Science.—South African
Journal of Science, xxiii (1926).

KIEw.
Société des Naturalistes—Mémoires, xxv, 2 (1917); xxvi, 1 (1917); xxvii, 1
(1926); Protocols (?), 1916-1917 (no date); Recueil des Travaux Scien-
tifiques, 1921-1922 (1924).

KURASHIKI,
Ohara Institute for Agricultural Research.—Berichte, iii, 2-3 (1926-1927).

La JOLLA.
Scripps Institution of Oceanography of the University of California.—Bulletin,
Technical Series, i, 1-9 (1927).

lxii. DONATIONS AND EXCHANGES.

La PuATA.
Museo de La Plata.—Anales, ii, 2 (1927); Obras Completas y Correspondencia
Cientifica de Florentino Ameghino, v (1926); Revista, xxx (1927).

LENINGRAD.

Académie des Sciences de VU.R.S.S.—Annuaire du Musée Zoologique, xviii,
1913, 4 (T.p. &c.), xxv, 1924 (1914-1925); xxvi, 1925, 3-4 (T.p.&c.) (1926);
Xxvii, 1926, 1-4 (1926-1927); xxviii, 1927, 1 (1927); Bulletin, Série vi, xix,
1925, 18 (Index) (1925); xx, 1926, 10-17 (1926-1927); xxi, 1927, 1-4 (1927);
“The Pacific-Russian Scientific Investigations” (1926); Travaux du Musée
Botanique, xix-xx (1926-1927).

Comité Géologique.—Bulletin, xxxii, 2-10 (T.p.&c.) (1913); xxxiii, 1-10
(T.p. &c.) (1914); xxxiv, 1-10, cover (1915); xxxv, 1-10 (1916); xxxvi, 1-10,
1917 (1917-1918); xxxvii, 1-10, 1918 (1919-1920); xxxviii, 1-10 (T.p.&c.),
1919 (1923-1925); xxxix, 1-10 (T.p.&c.), 1920 (1922-1924); xl, 1-10, 1921
(1922-1925); xli, 1-10, 1922 (1922-1926); xlii. 1-10 (T.p.&c.), 1923 (1923-
1925); xliii, 1-10, 1924 (1924-1926); xliv, 1-10, 1925 (1925-1926); xlv, 1-5, 1926
(1926-1927) ; xlvi, 1-8, 1927 (1927); Mémoires, xii, No. 1 (1915); N.S., Livr.
70, 77, 83-84, 87-88, 90-148, 150-154, 156-157, 159-160, 162, 167-168, 173-175
(1913-1927) ; Matériaux pour la Géologie Générale et Appliquée, Livr. 1-55
(1916-1926) ; 58-62 (1922-1924); 64-65 (1927, 1924); 68 (1927); 70 (1927);
80 (1924); 103-104 (1924, 1927); 111 (1927); 113-122 (1925-1926); Index
Général des Bulletins, 1882-1914 (1915); 1915 (1917); Bibliotheque
Géologique de la Russie, 1897-1900 (1901-1920); Travaux de la Direction
Centrale des Recherches Miniéres et de Prospection, Vol. i-iii (1922-1923) ;
Mineral Resources of the U.S.S.R., Parts 4, 12, 18, 20, 23, 24, 27, 28, 37
(1925-1927); Bulletins du: Comité Géologique; Section de Moscou, T. i, 1919
(1923); “Les Progreés de la Pétrographie en Russie”, by F. Loewinson-
Lessing (1923); “Bibliographie des Puits et des Sondages Artésiens en
Russie’, by S. Nikitin (1924); and 19 Separates (1913-1927).

Institut de Botanique Appliquée et dAmélioration des Plantes.—Bulletin of
Applied Botany and Plant Breeding, xvi, 1926, 5 (Index) (1926); xvii, 1927,
1-2 (1927); (continued as) Bulletin of Applied Botany, of Genetics and
Plant Breeding, xvii, 1927, 3 (1927); “The Agricultural Map of U.S.S.R.”,
edited by Prof. N. I. Vavilov (28th Supplement to the Bulletin of Applied
Botany and Plant Breeding) (1926); “The Best Varieties of the Spring
Wheats”, by V. V. Talanoff (1926).

Société Entomologique de Russie—Revue Russe d’Entomologie, xix, 1925, 3-4
(1926); xx, 1926, 1-4, 1926 (1926).

Société Russe de Minéralogie.—Matériaux pour la Géologie de Russie, xxvi, 1
(1918); Mémoires, 2me Série, li-liv (1918-1925).

LIEGE.

Société Géologique de Belgique.—Annales, xlviii, 4 (T.p.&c.) (1926); xlix, 1-3
(1926-1927); Bulletin (previously issued with Annales), 1, 1-6 (1926-1927) ;
Publications Relatives au Congo Belge et aux Régions Voisines, Année 1924-
1925, Annexe au xlviii, 1-2 (T.p.&c.) (1926); Année 1925-1926, Annexe au
xlix (complete) (1926).

Société Royale des Sciences de Liége-—Mémoires, 3me Série, xiii (1926).

DONATIONS AND EXCHANGES. 1xiii.

LIVERPOOL.
Liverpool School of Tropical Medicine-—Annals of Tropical Medicine and
Parasitology, xx, 4 (T.p.&c.) (1926); xxi, 1-3 (1927).

Lonpbon.

British Museum (Natural History).—A Monograph of the British Lichens,
Part ii, by Annie L. Smith (1926); Catalogue of Cainozoic Plants in the
Department of Geology, Vol. i. The Bembridge Flora, by Eleanor M. Reid
and Marjorie E. J. Chandler (1926); Descriptions of New Genera and
Species of Lepidoptera Phanaenae of the Subfamily Noctuinae (Noctuidae)
in the British Museum, by Sir George F. Hampson (1926); Economic

’ Leaflets No. 2 (1925); Economic Series. No. 44a (1925); Flora of Jamaica,
Vol. v, Dicotyledons, by W. Fawcett and A. B. Rendle (1926); Index
Animalium. Second Section. Parts vii-xi, by C. D. Sherborn (1925-1926) ;
Monograph of the Sea-snakes (Hydrophiidae), by Malcolm Smith (1926).

Entomological Society of London.—Proceedings, i, 1-3 (T.p.&c.) (1926-1927) ;
ii, 1 (1927); Transactions, Ixxiv, 1926, 1-2 (T.p.&c¢.) (1926); Ixxv, 1 (1927).

Geological Society—Geological Literature added to the Library during the
Years ended December 31st, 1924, and 1925 (1925-1926); Quarterly Journal,
lxxxii, 3-4 (T.p. & c.) (1926); Ixxxiii, 1-2 (1927).

Linnean Society—Journal, Botany, xlvii, 316-317 (1926-1927); Zoology, xxxvi,
244-246 (1926-1927); List, 1926-1927 (1926); Proceedings, 138th Session,
1925-1926 (1926).

Ministry of Agriculture and Fisheries.—Journal, xxxiii, 8-12 (T.p.&c.) (1926-
1927); xxxiv, 1-7 (1927).

Royal Botanic Gardens, Kew.—Bulletin of Miscellaneous Information, 1926
(1926); Hooker’s Icones Plantarum, Fifth Series, ii, 1 (1927).

Royal Microscopical Society.—Journal, 1926, 4 (T.p.&c.) (1926); 1927, Series
iii, xlvii, 1-3 (1927).

Royal Society.—Philosophical Transactions, Series B, ccxv, Nos. B 423-428
(1926-1927); Proceedings, Series B, c, No. B 705 (T.p. &c.) (1926); ci, Nos.
B 706-712 (T.p.&c.) (1927); cii, Nos. B 713-714 (1927).

Zoological Society.—Abstract of Proceedings, Nos. 279-291 (1926-1927); List of
Fellows, April 30th, 1926; Proceedings, 1926, 3-4 (T.p. &c. for pp. 683-1314)
(1926); 1927, 1; Special Notice, Jan., 1927 (1927); Transactions, xxii, 1-2
(1926-1927).

Los BANOS.
University of the Philippines: College of Agriculture-—The Philippine Agricul-
turist, xv, 6-10 (1926-1927); xvi, 1-5 (1927).

Lyon.
Société Linnéenne de Lyon.—Annales, Nouvelle Série, Ixxii, 1925 (1926).

MaDISoNn.
Wisconsin Academy of Sciences, Arts and Letters.—Transactions, xxi-xxii
(1924, 1926).

MapDRID.
Real Sociedad Espanola de Historia Natural.—Boletin, xxvi, 2, 8-10 (T.p. AY
(1926) ; xxvii, 1-7 (1927); Memorias, xiii, 2-3 (1926-1927).

lxiv. DONATIONS AND EXCHANGES.

MANCHESTER.
‘Conchological Society of Great Britain and Ireland.—Journal of Conchology,
xviii, 3-5 (1927).
Manchester Literary and Philosophical Society.—Memoirs and Proceedings, Ixx,
1925-26 (1926).

Manchester Museum.—Museum Publications 92-93 (1926).

Mania, P. I.
Bureau of Science of the Government ef the Philippine Islands.—Philippine
Journal of Science, xxxi, 2-4 (T.p.&c.) (1926); xxxii, 1-4 (T.p. &¢.);

xxxiii, 1-4 (T.p.&c.); Twenty-fourth Annual Report for the Year ending
December 31, 1925 (1927).

MELBOURNE.
Australasian Journal of Pharmacy, N.S. vii, 83-84 (Index) (1926); viii, 85-94
(1927). (From the Publisher.)
Commonwealth Bureau of Census and Statistics.—Official Year Book, Nos. 19-20,
1926-1927 (1926-1927).
Commonwealth Department of Health.Service Publication No. 32 (1926).

Commonwealth Department of Health, Division of Tropical Hygiene.—Service
Publication (Tropical Division) No. 9 (1926).

Council for Scientific and Industrial Research.—Bulletin Nos. 30-33 (1927);
Circulars 1, 3, 3, 4, 5, 7-10 (1919-1927); Journal, i, 1 (1927); Science and
Industry Endowment Fund—Notes on the Administration of the Fund
(Melbourne, 1927).

Department of Agriculture of Victoria.—Journal, xxiv, 11-12 (T.p.&c.) (1926);
xxv, 1-10 and Supplement (1927).

Department of Trade and Customs.—Fisheries. Biological Results of the
‘Fishing Experiments carried on by the F.I.S. “Endeavour”, 1909-14, v, 7
(T.p.&c.) (1926).

Field Naturalists’ Club of Victoria—The Victorian Naturalist, xliii, 8-12
(T.p. &e.) (1926-1927); xliv, 1-7 (1927).

National Herbarium.—“Census of Victorian Plants. Supplement No. 3”;
Supplement to Prof. Ewart’s “Weeds, Poison Plants and Naturalized Aliens
of Victoria’, by J. W. Audas and P. F. Morris (1925); “The Need for
Growing Softwoods for Paper Pulp and Paper Making”, by P. F. Morris
(From The Gum Tree, Sept., 1925); “A Valuable Swamp Grass’, by J. W.
Audas (From Journ. Dept. of Agric. Vict., June, 1925); Two Reprints by
J. W. Audas—‘The National Herbarium, Melbourne” and “The King’s Park,
Perth” (Vict. Nat., xliii, 1926); ‘Grasses of the Melbourne District’, by
P. F. Morris (From Vict. Nat., xliii, 1927).

National Museum.—Memoirs No. 7 (1927).

Public Library, Museums and National Gallery of Victoria.—Report of the
Trustees for 1926 (1927).

Royal Australasian Ornithologists’ Union—The Emu, xxvi, 3-4 (T.p. &c.)
(1927); xxvii, 1-2 (1927).

Royal Society of Victoria.—Proceedings, N.S. xxxix, 1 (1926).
University of Melbourne.—Calendar, 1927 (1926).

DONATIONS AND EXCHANGES. Ixy.

MExIco.
Instituto Geologico de Mesxico.—Anales, ii, 4-10 (1926-1927); Boletin, No. 43

(1924).

MILWAUKEE.
Public Museum.—Bulletin, iii, 2; vi, 3 (1927); Year Book, iv-vi, 1924-1926
(1926-1927).

Monaco.
L’Institut Oceanographique de Monaco.—Bulletin, Nos. 480-486 (T.p.&c. for

Nos. 469-486) (1926); 487-497 (1927).

MONTEVIDEO.
Museo de Historia Natural de Montevideo.—Anales, Ser. ii, ii, 2 (1926).

MONTREAL. :
Laboratoire de Botanique de VUniversité de Montreal.—Contributions, Nos. 6-8
(1926).
MUNCHEN.

Bayerische Akademie der Wissenschaften.—Abhandlungen, xxx, 9 and Map
(T.p.&c.) (1926); xxxi, 1-3 (1927); Sitzungsberichte, 1926, 1-3 (T.p. &c.)
(1926); 1927, 1 (1927).

NANTES.
Société des Sciences Naturelles de VOuest de la France.—Bulletin, 4 Sér., v,

1-4 (T.p.&c.), 1925 (1926).

NAPLES.
Stazione Zoologica di NapoliimPubblicazioni, vii, 3 (T.p.&c.) (1926); viii, 1

(1927).

New Haven, Conn.
Connecticut Academy of Arts and Sciences.—Transactions, xxviii, pp. 79-354

(1927).

New YorK.
American Geographical Society.—Geographical Review, T.p. & c. for xvi (1926) ;

xvii, 1-4 (1927).
American Museum of Natural History.—Bulletin, li, 1924-1925 (1924-1925); lv,
1926 (1926); Natural History, xxvi, 5-6 (T.p.&c.) (1926); xxvii, 1-3 (1927).
New York Academy of Sciences.—Transactions, iv (1884-1885).

OMSK.
Siberian Academy of Agriculture and Forestry.—Trudi (Transactions), ?
Supplement to iv, 1920-1925 (1925); v; vi, 1-9 (1926).

Orono, Maine.
Maine Agricultural Experiment Station.—Bulletins, 336, 337 (1927).

1xvi. DONATIONS AND EXCHANGES.

OSLo.

Det. Norske Videnskaps-Akademi i Oslo—Arbok, 1925-1926 (1926-1927) ;
Avhandlinger, I. Mat.-Naturv. Klasse, 1925-1926 (1926-1927); Register til
Forhandlinger og Skrifter utgit av Videnskapsselskapet i Kristiania, 1858-
1924 (1925); Skrifter, I. Mat.-Naturv. Klasse, 1925-1926 (1926-1927). —

Det Kongelige Frederiks Universitet.—Archiv for Mathematik og
Naturvidenskab, xxxix, 1-4 (T.p. &c.) (1925-1926).

OTTAWA.
Department of Agriculture.—Bulletin, N.S. No. 78; Pamphlet, N.S. Nos. 77, 82;
Report of the Director for the Year ending March 31, 1927 (1927); Report
of the Dominion Horticulturist for the Year 1925 (1926).

Department of Mines.—Report for the Fiscal Year ending March 31, 1926
(1926).

Department of Mines, Geological Survey of Canada.—Economiec Geology Series,
Nos. 2-3- (1926); Memoirs, 149-152 (1926-1927); Museum Bulletin, Nos. 438,
45 (1926); Summary Reports, 1925, pts. A-C (1926-1927); 1926, pt. B (1927).

Royal Society of Canada.—Proceedings and Transactions, Third Series, xx,
1926 (1926).

OXFORD. ~
Radcliffe Library, Oxford University Museum.—Catalogue of Books added
during the Year 1926 (1927). :

Pato ALTO.
Stanford University.—Bulletin of the Terrestrial Electric Observatory of

Dudley Herbarium, i, 1 (1927); Stanford University Publications, University
Series, Biological Sciences, ii, 4 (1923); iv, 1-3 (T.p.&c.) (1925-1926); vi, 1
(1927); Geological Sciences, i, 1 (1924).

ParRIs.
Journal de Conchyliologie, 1xx, 2-4 (1926-1927). (From the Publisher.)
Muséum National d’Histoire Naturelle—Archives, 6me Série, i; Bulletin, 1926,
1-6 (Ip. & ¢) (926); 1927, 1 (1927).
Société Entomologique de France.—Annales, xcv, 3-4 (T.p. & c.), 1926 (1926) ;
x¢vi, 1-2, 1927 (1927); Bulletin, 1926, 15-21 (1926); 1927, 1-2 and Supplement,
3-12 (1927).

PEKING.
Geological Survey of China.—Bulletin, No. 8 (1926).
Peking Society of Natural History.—Bulletin, Educational Series, No. 1 (1926) ;
“Flora Sinensis”’, Series A, i (1927).

PERM.

Institut des Recherches Biologiques a VUniversité de Perm.—Bulletin, i, 3-10
(Index) ; ii, 2-4, 6, 8, 9 and Supplement; iii, 2-10 (Index) and Supplement 2
(1923-1925); iv, 1-5, 10 (Index), Supplements 1, 2, 4 (1925-1926); v, 1-6
(1926-1927).

DONATIONS AND EXCHANGES. Ixvii.

PERTH.
Department -of Agriculture of Western Australia.—Journal, 2nd Series, ii, 4

(T.p. & c.) (1926); iv, 1-3 (1927).
Geological Survey of Western Australia.—Annual Progress Report for the
Year 1926 (1927); Bulletin, Nos. 84-87, 90, 91, 93 (1925-1927).

Government Statistician, Western Australia—Quarterly Statistical Abstract,
Nos. 243-246 (1926-1927).

Royal Society of Western Australia.—Journal, xii, 1925-1926 (1927).

PHILADELPHIA.
Academy of Natural Sciences.—Proceedings, ]xxviii, 1926; Year Book for the
Year ending December 31, 1926 (1927).

Academy of Natural Sciences, Entomological Section.—Entomological News
and Proceedings, xxxvii. 8-10 (T.p. & c.) (1926).

American Philosophical Society.—Proceedings, Ixv, 2-5 and Supplement to
No. 5 (separate index) (1926).

Wistar Institute of Anatomy and Biology—The Journal of Experimental
Zoology, xlvi, 2-4 (T.p. & c.) (1926-1927); xlvii, 1-3 (T.p. & c.); xlviii, 1-2
(T.p. & c.) (1927).

University of Pennsylvania.—Contributions from the Zoological Laboratory for
the Years 1924-25, xxiv (1925); Two Theses: “The Effect of Sulfur Dioxid
upon Plants: Methods of Study’, by L. A. Weierbach; “Chromosome
Structure and its Relation to the Chromosome Cycle, i’, by B. P. Kaufmann
(From Amer. Journ. Bot., xiii, 2, 1926).

Zoological Society of Philadelphia.—Fifty-fifth Annual Report of the Board
of Directors, 1927; Report of the Laboratory and Museum of Comparative
Pathology in conjunction with the Fifty-fifth Annual Report, 1927 (1927).

PITTSBURGH.
Carnegie Museum.—Annals, xvii (bound, complete), 1926-1927 (1927).

PLYMOUTH.
Marine Biological Association of the United Kingdom.—Journal, N.S., xiv, 3-4
(1927).

Porricl.
Laboratorio di Zoologia Generale e Agraria della R. Scuola Superiore
ad’ Agricoltura.—Bollettino, xix (1926).

PRAGUE.

Botanical Institute of the Charles University—Mykologia, i, 1-10 (T.p. & Ch)
(1924); ii, 1-5 (1925).

PRETORIA.
Botanical Survey of South Africa.—‘Diseases of Virginian Tobacco in South
Africa”, by Dr. E. S. Moore (Reprinted from Journ. Dept. Agric. South
Africa, Sept., 1926; Memoir, Nos. 7, 9 (1924, 1926).

1xviii. DONATIONS AND EXCHANGES.

PuUSA.

Agricultural Research Institute—Catalogue of Indian Insects, Parts 11-12
(1926-1927) ; Memoirs of the Department of Agriculture in India, Botanical
Series, xiv, 3, 5-6 (1927); Entomological Series, ix, 6-9 (1926); Review of
Agricultural Operations in India, 1925-1926 (1927); Scientific Reports, 1925-
1926 (1926).

RENNES.
Société Géologique et Minéralogique de Bretagne.—Bulletin, v, 2-4 (Index),
1924 (1926); vi, 2-4 (Index), 1925 (1927).

RicHMonpD, N.S.W.
Hawkesbury Agricultural College—H.A.C. Journal, xxiii, 8 (Index) (1926);
xxiv, 1-7 (1927).

Rio DE JANEIRO.
Instituto Brasileiro de Sciencias.—Boletim, Anno ii, 9-11 (1926-1927).
Instituto Oswaldo Cruz.—Memorias, 1926, xix, 1-2 (T.p.&c.) (1926); 1927, xx,
1 (1927).

RIVERSIDE, Cal.
Graduate School of Tropical Agriculture and Citrus Experiment Station
(College of Agriculture, University of California).—Papers Nos. 42, 51, 52,
89, 126-129, 131-140, 144-150, 153-157; T.p. & c. for iii, iv and v (1917-1927).

San DiIeEco.
San Diego Society of Natural History.—Annual Report for the Years 1923, 1924,
1925 and 1926 (1924-1927); “California Mammals’, by Frank Stephens
(San Diego, 1906); Transactions, i, 1, 3 (1905, 1911); ii, 1-3 (1914-1916) ;
iii, 1-4 (1917-1921); iv (complete) (1925); v, 1-6 (1927).

SAN FRANCISCO.

California Academy of Sciences.—Occasional Papers, xii; Proceedings, Fourth
Series, T.p. & c. for ii, pts. 1 and 2, 1913-1917, 1918-1926; T.p. & c. for xiv,
1925-1926 (1926); xv, 12-18 (T.p. & c.), 1926-1927 (1926-1927) ; xvi, 1-18, 1927
(1927).

Sao PAvLo.
Museu Paulista.—Revista, xiv-xv (1926-1927).

SARATOW.
Biologische Wolga-Station.—Arbeiten, viii, 4-5; ix, 1-2. (1926); Russische
Hydrobiologische Zeitschrift, i, 2-12 (T.p. & c.) (1922); ii-v (1923-1926) ;
veh, alert (Alby)

SEATTLE.
Puget Sound Biological Station of the University of Washington.—Publications,
Vv, pp. 109-179 (1926).

SENDAI.
Tohoku Imperial University.—Science Reports, 2nd Series, ix, 2 (1926); x, 2-3
(1926-1927); xi, 1 (1927); 4th Series, ii, 2-4 (T.p. & c.) (1926-1927); iii, 1
(1927).

DONATIONS AND EXCHANGES. lxix.

SHARON.
Cushman Laboratory for Foraminiferal Research.—Contributions, ii, 4; iii,
1-3 (1927).

St. Lovlis.
Missouri Botanical Garden.—Annals, vi, 4 (T.p, & c.) (1919); xiii, 3-4 (T.p. & c.)
(1926); xiv, 1-2 (1927).

STOCKHOLM.

Entomologiska Foreningen i Stockholm.—Entomologisk Tijdskrift, xlvii, 1926,
1-4 (T.p. & c.) (1926).

K. Svenska eviane tna eoniandemion.— ashe for Botanik, xx, 2-4 (T.p. & c.)
(1926); xxi, 1 (1927); Arkiv for Matematik, Astronomi och Fysik, xix, 3-4
(T.p. & c.) (1926-1927); Arkiv for Zoologi, xviii, 2-4 (T.p. & c.) (1926-
1927); Arsbok, 1926 (1926); Handlingar, Tredje Serien, ii, 1-8 (T.p. & c.)
(1925-1926) ; iii, 1-2 (T.p. & ce.) (1925-1926); Skrifter i Naturskyddsarenden,
Nos. 5-6 (1926).

SYDNEY.

Australasian Antarctic Expedition, 1911-14.—Scientific Reports, Series C, viii,
38 (1926). (From the Government Printer.)

Australasian Association for the Advancement of Science—Report of the
Seventeenth Meeting at Adelaide, August, 1924, xvii (1926).

Australian Museuwm.—Annual Reports of the Trustees for the Years ended 30th
June, 1924, 1925 and 1926 (1924, 1926); Australian Museum Magazine,
T.p. & c. for ii (1924-1926); iii, 1-4 (1927); Records, xv, 3-6 (T.p. & c.)
(1926-1927); xvi, 1 (1927).

Australian National Research Council.—Australian Science Abstracts, vi, 1-4
(1927).

Australian Veterinary Association.—Journal, Index to i (1926); ii, 4 (T.p. & c.)
(1926); iii, 1-3 (1927).

Botanic Gardens and Domains.—A Critical Revision of the Genus Eucalyptus,
vii, 7-8 (1926-1927), by J. H. Maiden, I.S.0., lately Government Botanist
and Director of the Botanic Gardens, Sydney.

Department of Agriculture, N.S.W.—Agricultural Gazette of N.S.W., xxxvii, 12
(1926); xxxviii, 1-11 (1927).

Department of Mines.—Annual Report for the Year 1926 (1927).

Department of Mines, Geological Survey of N.S.W.—Geological Map of the
Sydney District by T. L. Willan, B.Se. (1925).

Department of Public Health.Report of the Director-General for the Year
1925 (1927).

Education Department.—Education Gazette of New South Wales, xx, 12
(T.p. & c.) (1926-1927); xxi, 1-10 (1927).

Forestry Commission of New South Wales.—Annual Report for Year ended
30th June, 1926 (1926); The JASE PRSECEUEY Journal, ix, 10-12 (Index)
(1926); x, 1-9 (1927).

Government Statistician Quarterly Statistical Bulletin, Nos. 202-205 (Sept.,
1926-June, 1927) (1926-1927).

Institution of Surveyors—The Surveyor, xxxvi, 4-6 (1927).

Naturalists’ Society of New South Wales—The Australian Naturalist, v, 16
(1925); vi, 1-4 (1926-1927).

1xx. DONATIONS AND EXCHANGES.

Public Library of New South Wales.—Annual Report of the Trustees for the
Year ended 30th June, 1926 (1926).

Royal Society of New South Wales.—Journal and Proceedings, 1x, 1926 (1927).

Royal Zoological Society of New South Wales—‘A Monograph of the Australian
Loricates’’, by T. Iredale and A. F. B. Hull (Sydney, 1927); The Australian
Zoologist, iv, 5-6 (T.p. & c.) (1926-1927); v, 1 (1927).

The Medical Journal of Australia, 1926, ii, 22-26 (T.p. & c.) (1926); 1927, i,
1-26 (T.p. & ¢.) (1927); ii, 1-22 and Supplements 1-14 (1927); List of
Members of the B.M.A. in Australia (April, 1927). (From the Editor.)

TAIHOKU.
Government Research Institute, Department of Forestry.—Supplementa Iconum
Plantarum Formosanarum, Part iii (1927).

TASHKENT.
First Central Asiatic University. Bulletin, 13-15 (1926-1927).

TOKYO.
Imperial Fisheries Institute, Department of Agriculture and Forestry.—
Journal, xxii, 2-5 (1926-1927); xxiii, 1 (1927).
Imperial University of Tokyo.—Journal of the Faculty of Science, Section iii
(Botany), i, 2-8; ii, 1-2 (1927); Section iv (Zoology), i, 2-3 (1927).
National Research Council of Japan.—Japanese Journal of Botany, iii, 3 (1927).

TORONTO.
Royal Canadian Institute—Transactions, T.p. & c. for xv (1926); xvi, 1 (1927).

TOULOUSE.
Société d’Histoire Naturelle de Toulouse.—Bulletin, liv, 1926, 2-3 (T.p. & c.)

(1926); lv, 1926 (complete) (1926).

Trine, Herts.
Zoological Museum.—Novitates Zoologicae, xxxiii, 2-4 (T.p. & c.) (1926);
xxxiv, 1 (1927).

TRONDHJEM.
Det Kongelige Norske Videnskabers Selskab.—Arsberetning, 1925 (1926);
Skrifter, 1925-1926 and Statutter (1927).

TUNIS.
Institut Pasteur de Tunis.—Archives, xv, 4 (T.p. & c.) (1926); xvi, 1-3 (1927).

UPSALA.

University of Upsala—Bulletin of the Geological Institution, xx (1927);
Index to the Bulletin of the Geological Institution, xi-xx, 1912-1927 (1927);
“ober Evolutionistische Divergenz-Morphologie etc.’, by A. Soderstrom
(1927); “Zur Kenntnis der Zellarchitektonik, i’, by Adolf Soderstrom
(Upsala, 1926); Two Inaugural and one Academical Dissertations: ‘wher die
Hysteresis bei Glimmlichtentladungen mit Wechselstrom”, by Fredrik
Berglof (Uppsala, 1926); “tsber a -mercaptoketone und Verwandte Stoffe’’,
by Bertil Groth (Lund, 1926); Morphological and Biological Studies of the
Swedish Families of Hemiptera-Heteroptera, Pt. 1’, by Tore Ekblom
(Uppsala, 1926); Zoologiska Bidrag fran Uppsala, x-xi (1926-1927).

DONATIONS AND EXCHANGES. Ib:6:41,

URBANA.
American Microscopical Society.—Transactions, xlv, 4 (Index) (1926); xlvi,
1-4 (T.p. & c.) (Index) (1927).
University of Illinois.—Illinois Biological Monographs, x, 2-4 (T.p. & c.) (1926).

VIENNA.
Naturhistorische Museum in Wien.—Annalen, xl, 1926 (1926).
Zoologisch-botanische Gesellschaft in Wien.—Verhandlungen, |xxiv-lxxvi, 1924-
1926 (1926); Ixxvii, 1-2, 1927 (1927).

WARSAW.
Musée Polonais @Histoire Naturelle-—Annales Zoologici, v, 1-3 (1926).
Societas Botanica Poloniae.—Acta, iii, 1 (1925).

WASHINGTON.

Bureau of American Ethnology.—Bulletin 82-83 (1926-1927).

Carnegie Institution of Washington.—Publications, Nos. 342, 348, 373 (1924-
1926); Year Book, No. 25, 1925-1926 (1926).

National Academy of Sciences.—Proceedings, xii, 10-12 (T.p. & c.) (1926); xiii,
1 (Parts i-ii) (1926-1927), 2-9 (1927).

National Research Council.—Organization and Members, 1926-1927 (1926).

Smithsonian Institution—Annual Report for the Year ending June 30, 1925
(1926).

U.S. Coast and Geodetic Survey, Department of Commerce.—Serial Publication,
Nos. 350, 356, 366 (1926); Special Publications, Nos. 121, 123 (1926).

U.S. Department of Agriculture—Year Books, 1925, 1926 (1926, 1927); Bureau
of Entomology, Department Bulletins, Nos. 1380, 1426, 1428, 1429, 1439, 1453,
1469, 1472, 1476, 1482, 1487, 1490 (1926-1927); Department Circulars, 361,
395, 410-411 (1926-1927); Farmers’ Bulletins, Nos. 1494, 1495, 1503, 1531
1548 (1926-1927); Leaflet No. 2 (1927); Miscellaneous Circular, Nos. 70,
84, 104 (1926-1927); Technical Bulletin, Nos. 3-4 (1927); Report of the
Entomologist, 1925-1926 (1926); Twenty-nine Reprints from Journal of
Agricultural Research, xxxii, 11-12; xxxiii, 1, 2, 4-5, 7-9, 11-12 (1926); xxxiv,
4-9, 11 (1927).

U.S. Geological Survey. 47th Annual Report of the Director for the Fiscal
Year ended June 30, 1926 (1926); Bulletins, Nos. 782, 783 B-E (T.p. & c.), 784,
785 C-D (T.pp. & c.), 786 A-B (T.p. & c.), 788 A-D, 789, 790 A-B (T.p. & c.),
791, 792 A-B, 795 A-C (1926-1927); Professional Papers, 137, 139, 142 A-D,
147 A-C (T.p. & c.), 148 (1926-1927); Water Supply Papers, 530, 549-551,
553-555, 557, 559, 564, 565, 568, 575, 580 A-B (T.p. & c.), 583, 584, 592, 596 A-B
(1926-1927).

U.S. National Museum.—Bulletin, Nos. 100, Vol. ii, pt. 5; Vol. vi, pts. 2-3 (1926-
1927); 134-140 (1926-1927); Proceedings, T.p. & c. for lxviii (1927); lxix,
Arts. 1-23 (Nos. 2627-2649) (T.p. & c.) (1926-1927); Ixx, Arts. 1-23 (Nos.
2650-2672) (1926-1927); Ixxi, Arts. 1-23 (Nos. 2673-2695) (1927); Report
for the Year ended June 30, 1926 (1927).

WELLINGTON.
Department of Mines: New Zealand Geological Survey.—Bulletin, N.S., No. 29;
Palaeontological Bulletin, No. 12 (1927).

1xxil. DONATIONS AND EXCHANGES.

Department of Scientific and Industrial Research: Geological Survey Branch.—
Twenty-first Annual Report (New Series), 1926-1927 (1927).

Dominion Museum.—Bulletin, Nos. 6, 8 (1927, 1925); New Zealand Journal of
Science and Technology, viii, 5-6 (T.p. & c.) (1926); ix, 1-3 (1927).

New Zealand Institute —Transactions and Proceedings, lvii (1927); lviii, 1-2
(1927).

WELTEVREDEN.
Centraal Militair Geneeskundig Laboratorium.—Three Separates: ‘“Lop-

hoscelomyia annandalei var. djajasanensis nov. var.”, by S. L. Brug
(Genees. Tijd. voor Nederl.-Indie, |1xvi, 5, 1926); “Lophoscelomya annandalei
var. djajasanensis nov. var. (Diptera, Culicidae)”, by S. L. Brug (From
Bull. Soc. Path. Exotique, xix, 9, Nov., 1926); “Technische wenken voor het
verzamelen van muskieten’, by S. L. Brug (From Genees. Tijd. voor
Nederl.-Indie, Ixvii, 1, 1927).

Koninklijke Natuurkundige Vereeniging in Nederl.-Indie-—Natuurkundig
Tijdschrift voor Nederlandsch-Indie, Cover for xxxvi (1876); Ixxiv-lxxvi
(1915-1917) ; Ixxxvi, 3 (T.p. & c.); Ixxxvii, 1-3 (1927).

Woops Hote.
Marine Biological Laboratory.—Biological Bulletin, i-l] (1899-1926); li, 4-6
(T.p. & c.) (1926); lii, 1-6 (T.p. & c.) (1927); lili, 1-4 (1927); Zoological
Bulletin, i-ii (complete) (1897-1899).

PRIVATE Donors (and authors, unless otherwise stated).

Aneus & RoBerRTSON, Lrp., Sydney (donors).—“The Platypus’, by Harry Burrell,
C.M.Z.S. (Sydney, 1927).

AUSTRALIAN INSTITUTE OF TROPICAL MEDICINE, TOWNSVILLE, North Queensland
(donor).—One reprint: ‘Preliminary Note on the Transmission of the Hye
Worm of Australian Poultry”, by J. W. Fielding (From Aust. Journ. Exper.
Biol. and Med. Sci., iii, 1926).

Ciapp, F. G., New York City, U.S.A.—Two Reprints: ‘A Review of the Oil and
Gas Prospects of Australia” and “Oil and Gas Prospects in New Zealand”
(From Bull. Amer. Asscn. Petroleum Geologists, xi, 1, Jan., 1927, and x, 12,
Dec., 1926).

FERGUSON, Dr. E. W., Sydney (donor).—The British Medical Journal, Nos. 3419-
3473 (July 17th, 1926-July 30th, 1927).

GOVERNMENT OF INDIA CENTRAL PUBLICATION BRANCH, CALcuTTa, India (donor).—
Catalogue of Publications, compiled and corrected up to 31st December, 1925
(1926); 31st December, 1926 (1927).

INSTITUTION OF ENGINEERS, AUSTRALIA, SypNEY (donor).—‘Note upon the
Destruction of Wharf Piles by Crustacea and Mollusca in Australian Harbours’,
by A. C. Mackenzie (Paper No. 5, Vol. viii, 1927); Quarterly Bulletin of the
Institution of Engineers, Australia, iv, 14-16 (1927). ;

DONATIONS AND EXCHANGES. Ixxiii.

JANET, CHARLES, France.—“Constitution Orthobiontique des tres Vivants. i.
Théorie Orthobiontique” (Beauvais, 1925).

METROPOLITAN LIBRARY, PEKING, China (donor).—The China Foundation for the
Promotion of Education and Culture, First Report (March 1, 1926).

Meyrick, E., B.A., F.R.S., Marlborough, England.—Exotic Microlepidoptera, iii,
5-10 (1925-1926).

Musson, C. T., Sydney (donor).—‘“Carbon Assimilation by Plants’, by J. C. Bose
' (From “Scientia”, Sept., 1926); Presidential Address of Fourteenth Indian
Science Congress, by J. C. Bose (Lahore, 1927); Maine Agricultural Experi-
ment Station, Orono—Bulletins 313, 317 (19238, 1924); Department of Finance,
Imperial Plant Quarantine Service, Japan—Technical Bulletin No. 3
(Yokohama, 1925); Four Reprints by J. C. Bose from Proceedings of Royal
Society of London, Series B, Vols. 88 (1915); 89 (1916); 90 (1919); 93 (1922).

Sxvortzow, B. W., Harbin, China.—‘“Diatoms from Tientsin, North China”
(Reprinted from Journal of Botany, April, 1927).

Smiru, Dr. R. Greic, Sydney (donor).—Chemical Engineering and Mining Review,
xvii, 201, 1925); County of Northumberland, Education Committee: County
Agricultural Experiment Station, Cockle Park—Bulletin, Nos. 37-38 (1925-
1926); Dove Marine Laboratory, Cullercoats, Northumberland—Report for
Year ending June 30th, 1924 (1924); Royal Society of Tasmania—Papers
and Proceedings for Years 1924 and 1925 (1925-1926).

Steer, Mrs., Sydney (donor).—‘Monographie der Turbellarien. ii. Tricladida
terricola (Landplanarien)”. Text and Atlas. (2 vols.). By Dr. Ludwig von
Graff (Leipzig, 1899).

Tittyarp, R. J., M.A., D.Sc., F.R.S., Nelson, New Zealand (donor).—Journal of the
Royal Society of Arts, |lxxv, 3860, 1926; Five Reprints by Dr. Tillyard:
“Insect Control of Noxious Weeds” (From N.Z. Journ. of Agric., Feb., 1927);
“The Rhaetic ‘Crane-flies’ from South America not Diptera but Homoptera”’
(From Amer. Journ. Sci., xi, Mar., 1926); “A New Species of Spoon-winged
Lacewing (Family Nemopteridae, Order Neuroptera Planipennia) from
Western Australia” (From Journ. Roy. Soc. W. Aust., xii, pp. 1-4, 1925); “On
a Collection of Papuan Dragonflies, etc.” (From Rec. Aust. Mus., xv, 2, 1926);
“Upper Permian Insects of N.S.W. Part ii’ (From Proc. Linn. Soc. N.S.W.,
li, 3, 1926); ‘Notes on the N.Z. Wood-wasp Ophrynopus schauinslandi
Ashmead”, by E. S. Gourlay (From Trans. N.Z. Inst., 57, 1926).

Waters, A. W., F.L.S., F.G.S., Bournemouth, England.—‘“Ancestrula of Cheilosto-
matous Bryozoa, Pt. v’ (Reprinted from Ann. Mag. Nat. Hist., Ser. 9, xviii,
p. 424, 1926).

]xxiv.

LIST OF MEMBERS.

LIST OF MEMBERS, 1927.
ORDINARY MEMBERS.

1927 *Albert, Michel Francois, ‘““‘Boomerang’’, Elizabeth Bay, Sydney.

1905
1906
1922
1899

1927
1912

1913

1888
1925
1919

1907

1920
1923
1926
1912
1927
1912
1923
1921
1924
ALG yer
1920

1921
1910
1910
1926

1912
1899
1901
1927
1905
18930
1903
1899
1924
1920

1901
1920
1908

1300
1925

Allen, Edmund, c/o Chief Engineer for Railways, Brisbane, Q.

Anderson, Charles, M.A., D.Sc., Australian Museum, College Street, Sydney.

Anderson, Robert Henry, B.Sc.Agr., Botanic Gardens, Sydney.

Andrews, Ernest Clayton, B.A., F.G.S., Geological Survey, Department of Mines,
Sydney.

Armstrong, Jack Walter French, ‘“‘Callubri’, Nyngan, N.S.W.

Aurousseau, Marcel, B.Sc., c/o Mr. G. H. Aurousseau, ‘“Wondah”’, Bannerman
Street, Cremorne.

Badham, Charles, M.B., Ch.M., B.Sc., Bureau of Microbiology, 93 Macquarie Street,
Sydney. 5

Baker, Richard Thomas, The Crescent, Cheltenham.

Barnard, Colin, B.Sc., Commonwealth Research Station, Merbein, Vic.

Barnett, Marcus Stanley, c/o Colonial Sugar Refining Co., Ltd., O’Connell Street,
Sydney.

Benson, Professor William Noel, B.A., D.Se., F.G.S., University of Otago,
Dunedin, N.Z.

Blakely, William Faris, Botanic Gardens, Sydney.

Bone, Walter Henry, 15 Bond Street, Sydney.

Branch, Kenneth James Fergus, B.Sc., 99 North Steyne, Manly.

Breakwell, Ernest, B.A., B.Se., Agricultural High School, Yanco, N.S.W.’

Bredero, William Adrien Lewis, Box 127, Post Office, Orange, N.S.W.

Brewster, Miss Agnes, Girls’ High School, Sydney.

Brough, Patrick, M.A., B.Sc., B.Sc.Agr., ‘Kinross’, Billyard Avenue, Wahroonga.

Brown, Horace William, 871 Hay Street, Perth, W.A.

Brown, Miss Ida Alison, B.Sec., Geology Department, The University, Sydney.

Browne, William Rowan, D.Sc., Geology Department, The University, Sydney.

Burkitt, Professor Arthur Neville St. George Handcock, M.B., B.Sc., Medical School,
The University, Sydney.

Burns, Alexander Noble, ‘‘Lucerne’’, Lower Ferntree Gully, Victoria.

Burrell, Harry, 19 Doncaster Avenue, Kensington.

Burrell, Mrs. Harry, 19 Doncaster Avenue, Kensington.

Buzacott, James Hardie, Meringa (private bag), via Cairns, North Queensland.

Cadell, Miss Myall, ‘““Wotonga’’, Belgium Avenue, Roseville.

Cambage, Richard Hind, C.B.E., L.S., F.L.S., Park Road, Burwood.

Campbell, John Honeyford, M.B.E., Royal Mint, Ottawa, Canada.

Campbell, Thomas Graham, Australian Museum, College Street, Sydney.

Carne, Walter Mervyn, Government Botanist, Perth, W.A.

Carson, Duncan, c/o Winchcombe, Carson, Ltd., Bridge Street, Sydney.

Carter, Herbert James, B.A., F.E.S., “Garrawillah”’, Kintore Street, Wahroonga.

Cheel, Edwin, Botanic Gardens, Sydney.

Chisholm, Edwin Claud, M.B., Ch.M., Comboyne, N.S.W.

Clarke, Harry Flockton, c/o Colonial Sugar Refining Co., Ltd., Rarawai Mill,
Ba River, Fiji.

Cleland, Professor John Burton, M.D., Ch.M., The University, Adelaide, S.A.

Cooper, Mrs. A. G. S., B.Sc. (née Henry), Ogilvie Street, Denman, N.S.W.

Cotton, Professor Leo Arthur, M.A., D.Se., Geology Department, The University,
Sydney.

Crago, William Henry, M.D., 185 Macquarie Street, Sydney.

Cunningham, Gordon Herriot, Department of Agriculture, Biology Section, 71
Fairlie Terrace, Kelburn, Wellington, New Zealand.

* Life Member

LIST OF MEMBERS. 1xxv.

David, Sir Tannatt William Edgeworth, K.B.E., C.M.G., D.S.0., B.A., D.Sc., F.R.S.,
Sherbrook Road, Waitara.

de Beuzeville, Wilfred Alexander Watt, Tumut, N.S.W.

Dixson, Thomas Storie, M.B., Ch.M., 215 Macquarie Street, Sydney.

*Dixson, William, ‘‘Merridong’’, Gordon Road, Killara.

Dodd, Alan Parkhurst, Prickly Pear Laboratory, Sherwood, Brisbane, Q.

Drummond, Miss Heather Rutherford, ‘“Havilah’’, Glenbrook, N.S.W.

Dumigan, Edward Jarrett, Boys’ State School, Toowoomba North, Toowoomba,
Queensland.

Dwyer, Rt. Rev. Joseph Wilfrid, Bishop of Wagga, Wagga Wagga, N.S.W.

Enright, Walter John, B.A., West Maitland, N.S.W.

Flynn, Professor Theodore Thomson, D.Se., University of Tasmania, Hobart,
Tasmania.

Francis, William Douglas, Botanic Gardens, Brisbane, Queensland.

Friend, Norman Bartlett, 48 Pile Street, Dulwich Hill.

Froggatt, John Lewis, B.Sc., Department of Agriculture, Brisbane.

Froggatt, Walter Wilson, F.L.S., Young Street, Croydon.

Furst, Herbert Charles, 223 Macquarie Street, Sydney.

Garde, Miss Mary Lee, ‘‘Wavertree”, Kurraba Road, Neutral Bay.

Goldfinch, Gilbert M., ‘‘Lyndhurst’’, Salisbury Road, Rose Bay.

Grant, Robert, 24 Edward Street, Woollahra.

Gray, Archibald James, “Glendyne’’, Augusta Street, Concord.

Greenwood, William Frederick Neville, F.L.S., F.E.S., c/o Colonial Sugar Refining
Co., Ltd., Lautoka, Fiji.

Griffiths, Edward, B.Se., Department of Agriculture, 136 Lower George Street,
Sydney.

Gurney, William Butler, B.Sc., F.E.S., Department of Agriculture, George Street
North, Sydney.

Hacker, Henry, F.E.S., Queensland Museum, Bowen Park, Brisbane, Q.

Hale, Herbert Matthew, South Australian Museum, Adelaide, S.A.

Hall, Edwin Cuthbert, M.D., Ch.M., George Street, Parramatta.

Hall, Miss Lesley Damaris, B.Se., 8 Nicholson Street, Balmain East.

Hall, Leslie Lionel, ‘“Haldor’’, Drumalbyn Road, Bellevue Hill.

Halligan, Gerald H., F.G.S., ‘““Alameda’”’, Challis Avenue, Turramurra.

Hamilton, Alexander Greenlaw, “‘Tanandra’’, Hercules Street, Chatswood.

Hamilton, Arthur Andrew, “The Ferns’, 17 Thomas Street, Ashfield.

Hardwick, Frederick George, B.D.S., D.D.Se., Church and Macquarie Streets,
Parramatta.

Hardy, G. H. Hurlstone, The University, Brisbane, Q.

Harrison, Professor Launcelot, B.A., B.Se., Zoology Department, The University,
Sydney. :

Haviland, The Venerable Archdeacon F. E., St. Stephen’s Rectory, Portland, N.S.W.

Henry, Max, D.S.O., M.R.C.V.S., B.V.Se., Coram Cottage, Essex Street, Epping.

Hill, Gerald F., 5 Clifton Road, Hawthorn, Victoria.

Hopson, John, Jr., ‘“‘Dalkeith’’, Eccleston, N.S.W.

Hull, Arthur Francis Basset, Box 704, G.P.O., Sydney.

Hynes, Miss Sarah, B.A., ‘Isis’, Soudan Street, Randwick.

Irby, Llewellyn George, Forestry Department, Hobart, Tasmania.
Jackson, Sidney William, M.R.A.O.U., Belltrees, via Scone, N.S.W.
Jacobs, Ernest Godfried, ‘“Cambria’’, 106 Bland Street, Ashfield.

Johnston, Professor Thomas Harvey, M.A., D.Se., F.L.S., The University,
Adelaide, S.A. °
Kinghorn, James Roy, Australian Museum, College Street, Sydney.

Lawson, Augustus Albert, 9 Wilmot Street, Sydney.
Lea, Arthur M., F.E.S., 241 Young Street, Unley, Adelaide, S.A.

* Life Member.

LIST OF MEMBERS.

Le Plastrier, Miss Constance Emily Mary, ‘‘Carinyah’’, Provincial Road, Lindfield.

Le Souef, Albert Sherbourne, C.M.Z.S., Zoological Gardens, Taronga Park, Mosman.

Lindergren, Gustaf Mauritz, Secretary, Swedish Chamber of Commerce, 42 Bridge
Street, Sydney.

Lucas, Arthur Henry Shakespeare, M.A., B.Se., “Girrahween’’, William Street,
Roseville.

Mackerras, Ian Murray, M.B., Ch.M., B.Sc., Bureau of Microbiology, 93 Macquarie
Street, Sydney.

Mackinnon, Ewen, B.Sc., Council for Scientific and Industrial Research, 314
Albert Street, East Melbourne.

Mawson, Sir Douglas, Kt., D.Sec., B.E., F.R.S., The University, Adelaide, S.A.

McCarthy, Timothy, Bertram Street, Mortlake.

McDonnough, Thomas, L. S., ‘Iluka’, Hamilton Street, Randwick.

McHugh, Miss Mary Virgilius, St. Vincent’s College, Potts Point, Sydney.

McKeown, Keith Collingwood, Office of the Water Conservation and Irrigation
Commission, Leeton, N.S.W.

McKie, Rev. Ernest Norman, The Manse, Guyra, N.S.W.

McLuckie, John, M.A., D.Sc., Botany Department, The University, Sydney.

McNeill, Francis Alexander, Australian Museum, College Street, Sydney.

Mitchell, John, 10 High Street, Waratah, N.S.W.

Mitchell, Miss Dora Enid, B.Sc., Montague Street, Goulburn.

Moulden, Owen Meredith, M.B., B.S., ‘““Roma’’, Unley Road, Adelaide, S.A.

Mungomery, Reginald William, c/o Sugar Experiment Station, Bundaberg,
Queensland.

Murray, Patrick Desmond Fitzgerald, D.Sc., Zoology Department, The University,
Sydney.

Musgrave, Anthony, F.E.S., Australian Museum, College Street, Sydney.

Musson, Charles T., “Calala’”’, Nelson Road, Gordon.

Newman, Ivor Vickery, B.Sc., ‘‘Tip Tree’, Kingsland Road, Strathfield.

Newman, Leslie John William, F.E.S., ‘“Walthamstowe’”’, 5 Bernard Street,
Claremont, W.A.

Nicholson, Alexander John, M.Sc., F.E.S., Zoology Department, The University,
Sydney.

Noble, Robert Jackson, B.Se.Agr., Ph.D., c/o Mining Museum, George Street
North, Sydney.

North, David Sutherland, c/o Colonial Sugar Refining Co., Ltd., Broadwater
Mill, Richmond River, N.S.W.

Northcroft, Earle Fead, Biological Laboratory, Department of Agriculture,
Wellington, New Zealand.

O’Dwyer, Margaret Helena, B.Sc., Ph.D., Chemical Research Department, The
University, St. Andrews, Fife, Scotland.

Oke, Charles George, 56 Chaucer Street, St. Kilda, Victoria.

Oliver, Walter Reginald Brook, F.L.S., F.Z.S., Dominion Museum, Wellington, N.Z.

Osborn, Professor Theodore George Bentley, D.Sc., F.L.S., Department of Botany,
The University, Sydney. " .

Osborne, George Davenport, B.Se., Geology Department, The University, Sydney.

Perkins, Frederick Athol, B.Sc.Agr., Biology Department, University of Queensland,
Brisbane, Q.

Petrie, Arthur Hill Kelvin, M.Se., Department of Botany, University of Melbourne,
Carlton, Victoria.

Phillips, Montagu Austin, F.L.S., F.E.S., 57 St. George’s Square, London, S.W.,
England.

Pincombe, Torrington Hawke, B.A., Northwood Road, Northwood, N.S.W.

Pinkerton, Miss Ethel Corry, B.Sc., Ashford Street, Ashfield.

Priestley, Professor Henry, M.D., Ch.M., B.Se., Medical School, The University,
Sydney.

Pritchard, Denis Adrian, M.B., Ch.M., B.Sc., H.M.A.S. Moresby, G.P.O., Sydney.

Pulleine, Robert Henry, M.B., Ch.M., 163 North Terrace, Adelaide, S.A.

1927
1924

1925
1927

1919
1922
1916
1887
1909
1916
1926
1898
1923
1905

1911
1904

1926
1916

1904

1921
1902
1904

UG
1900

1909
1911

1926
1897

1927
1911
1924
1926
1922
1916
1916
1926
1923
1926
1903
1925
1910

1923

1923

LIST OF MEMBERS. ; xxvii.

Richardson, Keith Clifford, B.Se., Entomological Branch, Department of Agriculture,
136 Tower George Street, Sydney.

Roberts, Frederick Hugh Sherston, B.Sc., Prickly Pear Laboratory, Goondiwindi,
Queensland.

Roughley, Theodore Cleveland, Technological Museum, Harris Street, Sydney.

Rupp, Rev. Herman Montagu Rucker, The Rectory, Paterson, N.S.W.

Scammell, George Vance, B.Sc., ‘““Melrose’, 18 Middle Head Road, Mosman.

Shaw, Alfred Eland, M.R.C.S., L.R.C.P., F.E.S., 38 Harbour Street, Mosman.

Shiels, Mrs. N. L., M.Se., F.L.S. (née Collins), Norwood Avenue, Lindfield.

Sloane, Thomas G., F.E.S., Moorilla, Young, N.S.W.

Smith, G. P. Darnell, D.Se., F.I.C., F.C.S., Botanic Gardens, Sydney.

Smith, Miss Vera Irwin, B.Sc., F.L.S., “Cora Lynn’’, Point Road, Woolwich.

Stanley, George Arthur Vickers, B.Sc., ‘‘Clelands’’, Battery Street, Randwick.

Stead, David G., ‘‘Boongarre’’, Pacific Street, Watson’s Bay.

Steel, Miss Jessie Keeble, B.Sc., ‘“Helensburgh”, Marion Street, Killara.

Stokes, Edward Sutherland, M.B., Ch.M., Metropolitan Water, Sewerage and
Drainage Board, 341 Pitt Street, Sydney.

Sulman, Miss Florence, “Burrangong”, MecMahon’s Point.
Sussmilch, C. A., F.G.S., East Sydney Technical School, Darlinghurst, Sydney.

Taylor, Professor Thomas Griffith, D.Sc., The University, Sydney.
Tilley, Cecil Edgar, Ph.D., B.Sc., A.I.C., F.G.S., Sedgwick Museum, University
of Cambridge, Cambridge, England.

Tillyard, Robin John, M.A., D.Sc, F.R.S., F.L.S., F.E.S., C.M.Z.S., Cawthron
Institute, Nelson, New Zealand.

Troughton, Ellis Le Geyt, Australian Museum, College Street, Sydney.

Turner, A. Jefferis, M.D., F.E.S., Wickham Terrace, Brisbane, Q.

Turner, Rowland E., F.Z.S., F.E.S., The Needles Hotel, Port St. John’s, Pondo-
land, South Africa.

Veitch, Robert, B.Sc., F.E.S., Department of Agriculture, Brisbane, Queensland.

Walker, Commander John James, M.A., F.L.S., F.E.S., R.N., ‘“Aorangi’’, Lonsdale
Road, Summertown, Oxford, England.

Walkom, Arthur Bache, D.Sc., Macleay House, 16 College Street, Sydney.

Wardlaw, Henry Sloane Halcro, D.Sc., Physiology Department, The University,
Sydney.

Waterer, Arthur S., 14 Winchcombe Avenue, Haberfield.

*Waterhouse, G. Athol, D.Se., B.E., F.E.S., Macleay House, 16 College Street,
Sydney.

Waterhouse, Walter Lawry, B.Sc.Agr., ““Hazelmere’’, Chelmsford Avenue, Roseville.

Watt, Professor Robert Dickie, M.A., B.Sc., University of Sydney.

Wearne, Walter Loutit, ‘“Telarah’’, Collingwood Street, Drummoyne.

Weekes, Miss Hazel Claire, B.Sc., Zoology Department, The University, Sydney.

Welch, Marcus Baldwin, B.Sc., A.I.C., Technological Museum, Harris Street, Sydney.

Welch, William, F.R.G.S., ‘“Roto-iti’, Boyle Street, Mosman.

White, Cyril Tenison, F.L.S., Botanic Gardens, Brisbane, Q.

Whitley, Gilbert Percy, Australian Museum, College Street, Sydney.

Williams, Miss May Marston, M.Sc., “Bingera’’, 33 Day Street, Drummoyne.

Wood, Miss Lucy May, B.A., Zoology Department, The University, Sydney.

Woolnough, Walter George, D.Sc., F.G.S., Park Avenue, Gordon.

Wright, Fred, c/o Messrs. Elliott Bros., Ltd., O’Connell Street, Sydney.

Wymark, Frederick, 89 Castlereagh Street, Sydney.

HONORARY MEMBERS.
Hill, Professor J. P., Institute of Anatomy, University of London, University
College, Gower Street, London, W.C.1, England.
Wilson, Professor J. T., LL.D., M.B., Ch.M., F.R.S., Department of Anatomy, the
New Museums, Cambridge, England.

* Life Member.

xxviii. LIST OF MEMBERS.

1888
1902
1902

1902
1893

CORRESPONDING MEMBERS. -

Bale, W. M., F.R.M.S., 63 Walpole Street, Kew, Melbourne, Victoria.

Broom, Robert, M.D., D.Sc., F.R.S., Douglas, Cape Colony, South Africa.

McAlpine, D., Government Vegetable Pathologist, Department of Agriculture,
Melbourne, Victoria.

Meyrick, Edward, B.A., F.R.S., F.Z.S., Thornhanger, Marlborough, Wilts., England.

Spencer, Professor Sir Walter Baldwin, K.C.M.G., D.Se., F.R.S., The National
Museum, Melbourne, Victoria.

INDEX

(1927)

(a) GENERAL INDEX.

Aborigines, question of Royal Commission
of Inquiry, lii—acknowledgment of
resolution forwarded to Prime Minister,
liii.

Abstoma, species of, 242-3, 254.

Address, Presidential, i.

Albert, M. F., elected a member, xlix.

Alexander, C. P., The Interpretation of
the Radial Field of the Wing in the
Nematocerous Diptera, with special
Reference to the Tipulidae, 42.

Anderson, C., Note on the misapplication
of the name Thylacodes decussatus,
xlvi.

Andrews, E. C., congratulations to, iii.
Armstrong, J. W. F., elected a member,
xlv.

Asilidae, Further Notes on a new Classi-
fication of Australian, 387.

Australasian Association for the Advance-
ment of Science, i.

Australian Coleoptera, 222, 354—Diptera,
1, 33, 42, 284, 299, 336, 380, 387, 399—
Hesperiidae, 275—Proteaceae, the Xero-
phytic structure of the Leaf in, Part i,
258—marine algae, iv, 460—v, 555.

Balance Sheets, 1926, xxix-xxxi.

Barnard, C., Note on a Dicotyledonous
Fossil Wood from Ulladulla, N.S.W.,
113—congratulations to, liii.

Barrier Reef, Great, boring operations, ii.

Bees collecting bitumen, photograph of,
exhibited, li.

Beetle, fossil, exhibited, xlvii.

Bertram, B., elected a member, liii—
mosquito control in the Municipality of
Lane Cove, N.S.W., 563.

Brachycera, On the Phylogeny of some
Diptera, 380.

Bredero, W. A. L., elected a member, xliv.

Brough, P., Studies in the Goodeniaceae,
1, 471.

Brown, Ida A., Linnean Macleay Fellow
in Geology, appointed, 1927-28, x—re-
appointed, 1928-29, liii.

Butterflies, Australian, species exhibited
and lantern slides shown, li.

Cambage, R. H., elected a Vice-President,
xliv—congratulations to, iii—Note on
flight of flying squirrel, Petauroides
volans, XI|viii.

Campbell, T. G., elected a member, xliv.

Carter, H. J., Australian Coleoptera, Notes
and New Species, v, 222—elected a
Vice-President, xliv.

Cedar timber, exhibited, 1.

Chase, Eleanor E., obituary notice, iv.

Cheel, E., see Exhibits.

Cheilanthes vellea, The Anatomy of, 73.

Chert obtained from the Rhynie Chert
bed, Scotland, exhibited, xlix.

Chisholm, E. C., Additional Flora of the
Comboyne Plateau, 1926, 378.

Cleithrolepis granulatus, exhibited, liii.

Coleoptera, Australian, 222, 354.

Colloids, Influence of, upon Fermentation,
Ue

Comboyne Plateau,
the, 378.

Cooksey, D. F., notice of death, li.

Craft, F. A., elected a member, liii.

Additional Flora of

Cunningham, G. H., Gasteromycetes of
Australasia, vii. The Genera Disciseda
and Abstoma, 235—viii. The Genus
Mycenastrum, 245—ix. Keys to the

Genera and Species of the lLycoper-
daceae, 247.

Cylindro-conical stone from an aborigines’
camp, exhibited, xlv.

Deltopecten from the Illawarra District,
N.S.W., a new, 104.
Dendrobium, a new,
Queensland, 570.
Device for use in centering and mounting
microscopic objects, exhibited, xlviii.
Diptera, xvi, xxi—Australian, 1, 33, 42,
284, 299, 336, 380, 387, 399—Brachy-
cera, exhibited, xlviii.
Disciseda, species of, 236, 243, 254.
Diseases conveyed by Tabanidae, xx.
Dixson, Sir Hugh, obituary notice, iv.
Dixson, William, elected a member, xliv.
Donations and Exchanges, iii, xliii-xlvi,
xlix-l, lii, liii, lv.
Duke and Duchess of York,
XXVili.

for N.S.W., and

letter to,

1xxx.

Ecological Study of the Flora of Mount
Wilson. iv. Habitat Factors and Plant
Response, 161.

Elections, xxviii, xliv, xlv, xlix-l, lii-liii.

Elonichthys, exhibited, xlv.

Eremophila longifolia infested with rust-
fungus, exhibited, xlvii.

Estheria coghlani, exhibited, xlvii.

Estheriae of Australia, the Fossil, i, 105.

Exchange relations, iii.

Exhibits :—

Cheel, E., EHremophila
fested with
Hebenstreitia
corymbosa, liii.

Harrison, L., New species of Stratio-
drilus, liv.

Haviland, F. E., Device for use in
centering and mounting microscopic
objects, xlviii.

Hull, A. F. B., example of abnormal
Loricates, xlviii.

Lucas, A. H. S., specimens of English
primrose and cowslip and probable

longifolia, in-
rust-fungus, xlvii—
dentata and WSelago

hybrid, li.
Mackerras, I. M., series of Diptera
Brachycera, xlviii—series of Aus-

tralian Tachinidae, li.

McLuckie, J., specimens of chert from
the Rhynie Chert bed, Scotland, xlix.

Mitchell, J., cylindro-conical stone
from an aborigines’ camp, xlv—
fragment of fossil fish, Hlonichthys,
Xlv.

Musgrave, A., Papilio aegeus, remark-
able aberration of the female of, xlvii
—coloured drawing of Poecilopachys
bispinosa, liv.

Musson, C. T., Helix aspersa from
Gordon, Sydney, and England, xlv.

Nicholson, A. J., Papilio androgeus,
bilateral gynandromorph of, xlvii.
Pincombe, T. H., Cleithrolepis granu-
latus from Tambourine Bay, Sydney,

liii.

Stead, D. G., two extraordinary Malay
fishing lines, xlvii—remark on sharks
in Sydney Harbour, xlvi.

Waterhouse, G. A., Mosaic gynandro-
morphs of (a) Troides priamus
pronomus from Cape York (hb)

Papilio aegeus ormenus from Darnley
Is., also Papilio oberon from Santa
Cruz Is., xlvii—Hstheria coghlani
and a fossil beetle from Brookvale,
near Manly, xlvii—Ogyris oroetes,
Ogyris amaryllis and O. aenone, 1—
the new subspecies described in paper
(Aust. Hesperiidae, i), li.
Wearne, W. L., Cedar timber

portion of bed post embedded, 1.

with

INDEX.

Exhibits (continued) :—

Whitley, G. P., Iso rhothophilus
collected at Long Bay Baths, xlviii—
marine growth from off Long Reef,
N.S.W., li.

Ferguson, HE. W., collection of Amycterids
of, transferred to Macleay Museum, lii
—elected a Vice-President, xliv—Notice
of death, xlix—Presidential Address, i.

Fermentation, the influence of certain
colloids upon, 17.

Finckh, H. E., obituary notice, iv.

Fishing lines, two Malay, exhibited, xlvii.

Fleas and disease, xxv.

Fletcher, J. J., appreciation of, by Sir W.
Baldwin Spencer, xxxiii—memorial
lecture, ii—obituary notice, v.

Fletcher, the late J. J., and C. T. Musson,
On a case of natural hybridism in the
genus Grevillea, 122.

Flooding of the Hawkesbury River at
Windsor, the physiographic and
climatic factors controlling the, 133.

Flora, of Mount Wilson, an ecological
study of, pt. iv, 161—of the Comboyne
Plateau, additional, 378.

Fly breeding, prevention of, xxii.

Fossil, Estheriae of Australia, pt. i, 105
—wood from Ulladulla, N.S.W., 113.

Francis, W. D., elected a member, 1.

Garde, Mary L., elected a member, xliv.

Gasteromycetes of Australasia, vii, 235
—vili, 245—ix, 247.

Geology of the country between: Lamhb’s
Valley and the Paterson River, 85.

Greig-Smith, R., Macleay Bacteriologist
to the Society, notice of death, I—
influence of certain colloids upon

fermentation, iv, 17—v, 18—vi, 20—
summary of year’s work, ix. "

Grevillea, on a case of natural hybridism
in the genus, 122.

Hall, Lesley D., the physiographic and
climatic factors controlling the flooding
of the Hawkesbury River at Windsor,
11838},

Hamilton, A. G., elected a Vice-President,
1—lantern slides illustrating xerophytic
structure of leaf of Hakea, li—xero-
phytic structure of the leaf in Aus-
tralian Proteaceae, i, 258.

Hardy, G. H., further notes on a new
classification of Australian Robberflies
(Diptera, Asilidae), 387—on the phylo-
geny of some Diptera Brachycera, 380
—Notes on Australian and _ exotic
Sarcophagid flies, 447.

Harrison, L., elected President, xxviii—
see exhibits.

Haviland, F. E., see exhibits.

INDEX. 1xxxi.
Hawkesbury River at Windsor, the Mitchell, J., fossil Estheriae of Australia,
physiographic and climatic factors i, 105—new Deltopecten from the Illa-

controlling the flooding of the, 133.
Hebenstreitia dentata, exhibited, liii.
Hedley, C., obituary notice, vi.

Helix aspersa, exhibited, xlv.
Hesperiidae, Australian, i, 275.
Hitchcock, A. S., Two new species of

Setaria from Western Australia, 185.
Hull, A. F. B., congratulations to, iii—

elected a Vice-President, xliv—see

exhibits.
Hybrid between English primrose and

cowslip, exhibited, li.

Hybridism in the genus Grevillea, on a

case of natural, 122.

“Insects of Australia and New Zealand”,
Xi.
Iso rhothophilus, exhibited, xlviii.

Koalas, destruction of, li.
Kosciusko Plateau, the vegetation of the,
1, LSA.

Lamb’s Valley, geology of the country
between, and the Paterson River, 85.
Lawson, Prof. A. A., notice of death,
XXViii.

Lea, A. M., descriptions of new species of
Australian Coleoptera, xix, 354.

Life Membership Subscription, xliii, xliv.

Linnean Hall at Elizabeth Bay vacated,
xlv.

Linnean Macleay Fellowships,
tions invited, li, lii.

Liphyra brassolis, lantern slide of pupa
of, exhibited, liii.

Liversidge, Prof. A., notice of death, li.

Lizards, a notice on _ reproductive
phenomena in some, 25.

Loricates, abnormal, exhibited, xlviii.

Lucas, A. H. S., Notes on Marine Algae,
iv, 460—v, 555—see Exhibits.

applica-

Mackerras, I. M., Linnean Macleay Fellow
in Zoology, Notes on Australian Mos-
quitoes (Diptera, Culicidae), i, 33—1ii,
284—reappointed, 1927-28, x—summary
of year’s work, x—see Exhibits.

Malaria, xv.

Malloch, J. R., Notes
Diptera, x, 1—xi, 299—xii,
399.

Marine growth, exhibited, li.

McHugh, Mary V., elected a member, xlix.

McKie, Rev. E. N., elected a member, lii.

McLuckie, J.. and A. H. K. Petrie, an
ecological study of the flora of Mount
Wilson, iv, 161—the vegetation of the
Kosciusko Plateau, i, 187—see Exhibits.

Medical and Veterinary Entomology in
Australia, Xv.

336—xiili,

on Australian «

warra 104—-see
Exhibits.

Mosquito control in the Municipality of
Lane Cove, N.S.W., 563.

Mosquitoes, notes on Australian, i, 33—ii,
284.

Moulton, D., new gall-forming Thysan-
optera of Australia, 153.

Mount Wilson, an ecological study of the
flora of, iv, 161.

Murray, P. D. F., congratulations to, iii.

Musson, C. T., see Exhibits.

Musson, C. T., and the late J. J. Fletcher,
on a case of natural hybridism in the
genus Grevillea, 122.

District, N.S.W.,

Native bears, destruction of, li.

Nematocerous Diptera, the interpretation
of the radial field of the wing in the,
42.

Nicholson, A. J., elected to Council, 1—
see Exhibits.

Obituary Notices, Eleanor E. Chase, iv—
Sir Hugh Dixson, iv—H. E. Finckh, iv
—J. J. Fletcher, v, xxxiii—C. Hedley, vi
—E. H. Rennie, ii.

O’Dwyer, Margaret H., congratulations to,
iii.

Ogyris oroetes, O.
aenone, exhibited, 1.

Oke, C. G., elected a member, xlv.

Osborn, T. G. B., elected a member, lili.

Osborne, G. D., the geology of the
country between Lamb’s Valley and the
Paterson River, 85.

amaryllis and O.

Pacific Science Association, formation of,
ii.

Pan-Pacific Congress, Third, ii.

Papilio aegeus, aberrant, exhibited, xlvii
—P. aegeus ormenus, aberration, ex-
hibited, xlvii—P. androgeus, exhibited,
xlvii—P. oberon, exhibited, xlvii.

Paterson River, geology of the country
between Lamb’s Valley and the, 85.
Petauroides volans, note on flight of,

xviii.
Petrie, A. H. K., congratulations to, xlix
—see McLuckie, J., and A. H. K. Petrie.
Phalacrocorax carbo, large black shag,
note on feeding habits of, lii.
Physiographie and climatic factors con-
trolling the flooding of the Hawkes-
bury River at Windsor, 133.
Pincombe, T. H., see Exhibits.
Placentation in some lizards,
xliii.
Poecilopachys bispinosa,
ing of, exhibited, liv.
Presidential Address, i.

note on,

coloured draw-

Ixxxii.

Prize offered by Société de Physique et
d’Histoire Naturelle de Genéve, 1.

Proteaceae, Australian, xerophytic struc-
ture of the leaf in, i, 258—lantern slides
shown, li.

Radial field of the wing in the Nemato-
cerous Diptera, the interpretation of,
42.

Rennie, Prof. E. H., reference to death,
ii.

Reproductive phenomena in some lizards,
a note on, 25.

Richardson, K.C., elected a member, xliv.

Robberflies, further notes on a new classi-
fication of Australian, 387.

Rupp, Rev. H. M. R., elected a member,
xlix—a new Dendrobium for N.S.W.,
and Queensland, 570.

Sarcophagid flies, notes on Australian
and Exotic, 447.

Science Association, Pacific, formation of,
ii.

Scincid lizard, Lygosoma _ (Hinulia)
quoyi, Placentation and other phe-
nomena in, 499.

Selago corymbosa, exhibited, liii. '

Selby, Doris A., elected a member, liii.

Setaria, two new species of, from Western

_ Australia, 185.

Sharks, note on occurrence of, xlvi.

Stead, D. G., destruction of native bears,
li—note on occurrence of sharks, xlvi
—note on feeding habits of large Black
Shag, Phalacrocorax carbo, lii—note on
wafer worm, Leptoplana australis, liii
—photograph showing bees collecting
bitumen, li—see Exhibits.

Stratiodrilus, new species of, exhibited,
liv.

Studies in the Goodeniaceae, i, 471.

Sussmilch, C. A., congratulations to, liii.

Tachinidae, Australian, exhibited, li.

Thylacodes, note on, xlVvi.

Thysanoptera of Australia,
forming, 153.

new gall-

INDEX.

Tillyard, R. J., congratulations to, liii—

notice of book by, xi.

Tipulidae, the interpretation of the radial
field of the wing in the Nematocerous
Diptera with special reference to the,
42.

Troides priamus pronomus,
exhibited, xlvii.

aberration,

Vegetation of the Kosciusko Plateau, i,
187.
Veterinary Entomology in Australia, xv.

Wafer worm, Leptoplana australis, note
on, liii.

Walkom, A. B., congratulations to, iii—
granted leave of absence, iii—greetings
from, i—welcome extended to, liii.

Waterhouse, G. A., appointed Acting Sec-
retary, iii—Australian Hesperiidae, i,
275—coloured lantern slides of butter-
flies shown, li—elected Hon. Treasurer,
xliv—lantern slides of pupa of Liphyra
brassolis shown, liii—see Exhibits.

Waterhouse, W. L., elected a member,
xlix.

Wearne, W. L., see Exhibits.

Weekes, H. Claire, Linnean Macleay

Fellow in Zoology, appointed, 1927-28,
x-xi—reappointed, 1928-29, liii—note on
reproductive phenomena in some
lizards, 25—note on placentation in
some lizards, xliii—Placentation and
other phenomena in the Scincid Lizard,
Lygosoma (Hinulia) quoyi, 499.

Western Australia, two new species of
Setaria from, 185.

Whitelegge, T., notice of death, 1.

Whitley, G. P., see Exhibits.

Williams, May M., Linnean Macleay
Fellow in Botany, reappointed, 1927-
28, x—summary of year’s work, x—

the anatomy of Cheilanthes vellea, 73.
Wood, dicotyledonous fossil, from Ulla-
dulla, N.S.W., 122.
Woolnough, W. G.,
xliv.

congratulations to,

(b) BIOLOGICAL INDEX.

Synonyms are printed in italics; new names in SMALL CAPITALS.

Abstoma 235, 241, 253-4 Acacia penninervis.. 196,198 Aciphylla glacialis .. 217
purpureum .. 243, 254, 256 rubida . 196 Ackmana ate TEL
RETICULATUM.. . 242, 254 siculiformis . 198 Acracantha ROME ne 0) | 5s)

Acacia .. 271 Acaena sanguisorbae .. Acutipula omissinervis .. 59
alpina ng, Js 194-5,197-8,217 Adelodus LEGS Ee OO
decurrens .. 198 Acantholimnophila 50 eAdelphomvilaiel alee OU
doratoxylon .. 155 Acanthophora orientalis 555 Adenanthos cygnorum 258
neriifolia 7 LS Acer , . 114 ? Adiochosia hyalipennis 308

. 284, 286

Aedes .
aculeatus — 285-6, 294-5
aegypti EAE EBX:
albirostris . 285, 288
alboannulatus . 290, 565
andersoni 287, 289, 291-2
antipodeus . 285, 288
australis 292-3
brugi : e288
burpengaryensis ath

285, 287, 289, 293- 4

camptorhynchus ee
285-6, 289, 290-1
clelandi .. . 285, 287, 294
concolor BN echt CMe OD) OL)
cunabulanus 287, 289, 292

flavifrons Ee TS
285-7, 289, 290, 565
imprimens . 285, 288
luteifemur 286-7, 293
MACLEAYANUS . 287, 291
nigrithorax 287, 289, 292-4
nivalis 285, 287-8, 292-3
normanensis 285, 287, 294-6
notoscriptus ». 565
occidentalis Pe . 290
purpureiventris . 287, 294
queenslandis . 298, 565
rubithorax 285-6, 288, 297
sagax . 285, 287, 289, 292
stricklandi 285-7, 290
theobaldi 285-7, 294-7

theobaldi form EIDS-
VOLDENSIS . 287, 295
vandema . 289, 290
variegatus Lil iuiEtye Hone ©.<
vexans . 288

vigilax 284-8, 290, 296-7, 565

vittiger 285-6, 289, 292-3
wilsoni 285, 292-3, 295
Aedimorphus . 286
alboscutellatus 5 ASU
vexans Sh . 288
Agamofilaria tabanicola xx
Agromyza . . 424, 426
ALBISQUAMA . 425
citreifemorata . 428
flavocentralis . 427
lantanae .. 426
PALLIDICENTRALIS . 427
phaseoli 425-6
tricolor 5 Le
TRISPINA : 9 425
Aldrovandia PE oan ieee eS BG)
caine fs bs Jae Mee bY)
Allecula morio 3 BS
Alnus . 114

Alsophila 165, 167, 172, 182, 184
Alysia manducator .. XXVii
Amalopina dicranotoides 48
elegantula .. 48
sibiriensis Beene al a thas C45)
Amenia
DUBITALIS Mininae, eee noe

INDEX.
Amenia imperialis .. 342-3
leonina 342-4
parva 342-4
Amphibolia . 349
Amphibolurus barbatus
Xi, 532
Amphichroum sp. .. .. 232
Amphicyphus . 399, 400
reticulatus .. 400
Amphilimnobia leucopeza 68-9
Amphineurus .. .. .. 56
Amsonia 3 5 . 484
Anadyomene Brownii 5 BS
Muelleri 56.
Anagallis arvensis .. 194-5
Anastellorhina bicolor .. 335
Anausis metallescens. .. 230
Anaxo CYLINDRICOLLIS .. 233
eylindricus 5 ABR}
dentipes 233

Andersonia tasmanien sis 291

Anguillaria dioica . .. 570
Anilara En 5.) 220
anthaxoides .. . 223
Anisynta sphenosema 6 AUT
tasmanica : 5 Att
tillyardi 277
Anopheles .. 33- 4, 36
amictus Xviii, 33-7
annulipes

xviii, 33-5 -5, "37, 39- 41, 565
atratipes A) BRE, Orso Gal
bancrofti xviii, 34-5, 38-40
corethroides .. 33, 40
corethroides .... .. 40
maculipennis xviii
mastersi 33, 35
punctulatus xviii, 33-4
stigmaticus 33-5, 37-40
Anoplura XXVii
Anthotium ait we yee: af ica
ANTHRACOMYIA 303, 318-9
ATRATULA so lly)
Antithamnion .. .. 460
Antocha 63, 68
saxicola .. .. 68-9
Aphesia .. 394
Aphrophila .. 54-5
APINOCIS 56 axtfal
VARIIPENNIS 371-2
Apterygota SUR ia ato. o ut
Aralia cA . 114
Architipula Rene Shyu ft iL
Armeria ; a .. 484
Arthrophycus . 560

Arthropodium eG
tum ay 197-8
Arthropteris obliterata a5 Ble
Asilus : 388, 392-3
crabroniformis 392-3

Asplenium flabellifolium
95,198

Astacoides madagascari-
ensis ap oe Whey Ao mnie SOL

1xxxili.

Astacopsis serratus oe Liliv
CASINLAMI CUS Mums eee em nL LN
Asteia oes . 445
Astelia alpina .. 4 AY
Astelobia . 54-5
Aster novae-anglicae . 489
Atarba av 52, 63
picticornis . 52-3
Athemistus e234
albofasciatus + 204
conifer . 234
Atherosperma . 175-6
moschatum 5 alee!
Atomasia en P OO:
Australina . 400, 421
geniseta so CYA
Austrolimnobia . 54-5
Avrainvillea clavatiramea 557
erecta 5 DHS
papuana . 5 DDO
Azorella cuneifolia a. 5 paalts
dichopetala 217-8
Bacillus levaniformans.. 20
vulgatus ATELAAT = OMERS TRY S OMI

Baeckea Gunniana ..
196, 205- 6, 208- 9

Banksia . 173, 258, 260
integrifolia 259
marginata 56 (PADS)
serrata no dberey
spinulosa So Abtttss

Banksinella no CRS
brugi : 1. 288
lineatopennis. a RSH

Barbarea vulgaris .. .. 217

Baris aie 370- 1
albigutta 365-6
albopicta . 365, 368
angophorae .. 362, 365, 368
apicinivea tO.
australiae . 361, 364
BARRONENSIS 5 OY)
basirostris st moun 64
BITUBERCULATA . 358, 365
bryanti Sa AO
CAIRNSENSIS eos
COELESTIS 50. ati
CYANEOTINCTA es . 363
DILATATIFRONS Fone Le oe OLIN
ebenina = O09
elliptica . 364
EPISTERNALIS ooo
GERALDTONENSIS OOS
glabra . 365
glabra Las OOS
HOPLOCNEMIS . 359, 369
TLLEPIDA .. 362
LATERICOLLIS so HOS
MELANOCHROA S6 ON
MELANOSTETHA 5 hol
microscopica ae . 368
NIVEODISPERSA .. 358, 365
niveonotata . 365, 368
oblonga .. Sakae

1xxxXiv.

Baris oblonga .. ee OS:
ORTHODOXA no BHOK))
PARVONIGRA . 360
PRAEMORSA Bie 365
PULCHRIPARVA i 359, 360
PULCHRIPENNIS .. 367, 369
SETIPENNIS . 364
SETISTRIATA .. .. 360
sororia mi 362, 365, 368
subopaca x0 aH)
transversicollis SOS
TRISINUATA Pe Ole
vagans 360-1, 363
VULNERATA . 363

Bathypogon 2 395-6

Batrachomyia . 435, 440
atricornis . 440
flavicornis . 440
MAJOR 440-1
STRIGIPES aS 440-1

Batrachus dubius .... lii

Benjaminella . 435, 440
albifacies so oe 25

Blechnum capense .. 162,181
penna-marina ae 194-5

Blepharotes . 3890, 393
abdominalis .. 393
brisbanensis oo ays}
corarius 56 GH)3}
flavus 65) BR}
punctatus 5 as}
vivax 3 393

Boodlea coacta’ - i555: 6

Bossiaea foliosa 195-6, 198

Botanobia 435, 444-5

Bovista 253-4
anomala 5 ABS
argillacea . 255
aspera 5 ZB
brunnea 254-5
candida .. . 255
cervina a 255
circumscissa S255
debreciensis 2 12Dd
gigantea 5 PASS)
hyalothrix q ABD
hypogaea ~ 255
lilacina 5255
muelleri . 255
olivacea .. 255
ovalispora _ Pay)
purpurea . 254
pusilla - 255
suberosa . 255
subterranea 51255
tomentosa 2 255
tunicata 5 Oy,

Bovistella 253-4
aspera . 255
australiana te en
bovistoides . 254, 257
coprophila n 3 Pat
glabrescens > 205
Gunnii . 205

INDEX.
Bovistella nigrica .. -255
pusilla aera
rosea 50. ASD
scabra . 255
verrucosa 254
Brachycome cardiocarpa
var. alpina 217-8
decipiens : mir eesee lO
scapiformis 195, 197-8, 217
Brachypremna on 58
candida .. ao Bee 9
Be Cacia ‘ 388, 394
bella 5 5 Oo
fenestrata .. 394
fulva SOO:
maculata co etl
pulchella .. 394
ruficornis 50 Oe!
victoriae Brn gated . 394
Brunella vulgaris ou od)
Brunonia 5 5 Rae er
Bryopsis plumosa Hel eek BR x
Bulbine bulbosa 50 Ye
Bursaria spinosa . 160
Cabasa so oe!
rufithorax .. o94
Caiusa 5 0 AeA
surcoufi . 5b) BR
Calamites .. .. 90-1
Calendula . . 484
Caligus xlvili
Callicoma ..
114, 120, 162, 176, 181- 2
serratifolia 175-6, 181
Calliphora ee nee 300-3
ACCEPTA .. 307, 311, 314, 316
APICALIS 306, 312-3
ASSIMILIS 5 OS Bil7
augur 306, 309-11
AURIVENTRIS = 30K. a5
AUSTRALICA . 806, 314
CENTRALIS 5 5 SG, aBlal
CLARKI 5 atl, silils Balko
dispar . 306, 312
elliptica 5 Bets)
erythrocephala ..
303, 306, 312-4, 319
fulviceps ee . 302
FUSCOFEMORATA 306, 309-11
hilli . 306, 309
lateralis : 55. BILD)
MACLEAYI 306, 310-1
METALLICA 5 BOM, Bally
MINOR . 007, 314
oceaniae Sp eee ly)
ochracea OOD 08
ochracea form NIGRI-
THORAX 56 oo GU, CLE
PLEBEIA 5 De Ass, B37
pubescens - 312
ROBUSTA 306, 313- 4
rufiventris » oly)

Suri. Soden ate

Calliphora stygia ..
XXiv, 305, 308- 9

tessellata u oS
tibialis 305, 308, 315-6
villosa AN eee oly)
XANTHOCERA .. . 306, 313

Callistemon Sieberi 194-5,
198-9, 204, 206, 208-9, 211-2
Callithamnion 460-463, 467, 469

angustatum . Abt
angustatum Sak Pasa O
brounianum .. 461, 464
brounianum . 464
conspicuum ‘ 460-1
crispulum .. 461, 469
dasytrichum 55 Atel
dasyurum . 461
dasyurum ego LOG
debile . 461, 469
formosum Me ecg UI |
hirtum .. 461, 465
latissimum . 461
latissimum 56° 400
paradoxum ~ Aueil
paradozum .. . 466
plumigerum .. 460-1
scopula . 461
scopula . 468
stuposum . 461
superbiens 460-1
tingens 460-1
violaceum . 461
violaceum . 468
wollastonianum 5 AuGal
wollastonianum . 465
Calogyne 3 é we AW
Calolimnophila Se SETS
Caltha introloba 2lGer2ies
Calvatia 5 YS
aurea Pectin Sear a BG
caelata 253, 255-7
candida .. 253, 255-7
candida var. rubro-
flava 256-7
cyathiformis . 256
favosum . 256
Fontanesti . 256
gigantea 253, 255-7
lilacina 253, 256-7
maxima .. : 256
occidentalis bo PADD
olivacea .. ao SMO
primitiva ao ADD
rubro-flava po 7A
sinclairii .. 256
Campanula Hae OA:
Campbellomyia ia) ORO
Campolene CYANEA 5 BVA
nitida P Bie ABA
Carcharias arenarius 5 xdhial
Carcharinus brachyurus xlvi
Cardiothorax australis
var. , PANE
consimilis Nido

Carex breviculmis ..
Gaudichaudiana
Cassinia aculeata

217-8
. 212

192, 194, 196, 198-9

Castiarina DEUQUETI a 225
DILATATA 226-7
FLAVO-VIRIDIS bo PAT
FOSSORIA Per a20

Casuarina .. aerate!

Catastoma 66 De
anomalum .. 256
circumscissa sot 2D
debreciensis .. a6 CANS
hyalothrix o. 256
hypogaea ce ZS
Muelleri s5 D8
pedicellata .. 256
purpurea ss CASO)
subterranea S256

Catosperma 472

Caulerpa Xe 2 BRR, 558 9
abies So BN)
Brownii.. 6 DY)
cactoides So oo BY)
clavifera . 5dd, 559
cupressoides 556
fastigiata Se DDS
flexilis co. HRY)
Hedleyi Pe 6a)
hypnoides 56 BD)
laetevirens 56 ERG
ligulata eS
Muelleri so BBL)
Pickeringii 50 DY)
sedoides so DOD
trifaria 56 Say
tristicha 55 DD
Webbiana form _ tor-

mentella SF 559

Caviceps : 435, 442

flavipes . 442

Celmisia longifolia ..

195, 215, 217-8

Ceramium clavulatum .. 555
Ceratocheilus 57, 69
australasiae . 68-9
edwardsi . 68-9
flavirostris 36 to 6. ised)
Ceratolauxania . 400, 408
TASMANIENSIS .. 408
tetanocerina 408-9
Ceratolimnobia 54-5, 62
Ceratopetalum .. 114, 120,
161, 167-8, 172-4, 176, 184
apetalum priors awa LMS)
Ceratophyllus fasciatus xxvi
Cerdistus 387, 390-3
fulvipubescens 392
Cerodonta 423
australis 423
robusta 423
VITTIGERA 423
Cerozodia 50
Chaetogaster 3i5)33

INDEX.

Chaetogaster violacea .. 353
Chalcides ocellatus ..
xii, 27, 32, 500
tridactylus xliii, xliv,
25, 27-8, 30, 32, 499, 500,
503, 506-7, 537, 546-550, 553
Chalecoponera metallica xxvii

Chamaclaucium uncin-
atum 5 aid

Chantransia as . 460

Cheilanthes | TRS ts TSN 803
emGllerrn os ov (eh tis 0 1
gracillima aor o
lanuginosa .. Ts TH, TD)
lendigera (35
microphylla Ua, U5)
persica U5 5 US)

tenuifolia 73-5, 77, 79, 81, 83

vellea x, 73-5, 77, 79, 83-4
Chiloglottis Gunnii 3) LOS
Chionea = su tete boys
Chloromerus » 430

GRACILIS 431-2

MACULIFEMUR 431-2

NIGRIFEMUR 431-2

pallidior AL ASE A Silt

purus E 431433

purus form MACU-

LIFERA . 431, 433

purus form purus 431, 433
purus form vaRIANs 431, 433

TRIM ACULATA esl, 48333
Chloropisca .. 429
MONTICOLA . 430
notata 429-30
‘SUBNOTATA 429-30
Chlorops Badia a.
Chlororhinia 1 329.7332
viridis bo Oe
Chlorotachina 1 BDS
flaviceps Sade
CHOLEOTHRIPS 5 Aly)
GEIJERAE 5 155-6
Chondria 555, 560-1
caerulescens . 560, 562
curdieana 562
dasyphylla e560
fusifolia 50 BOAL
IRIDESCENS 50 Boll
RAINFORDI . 560
sedifolia 560
tenuissima BaD 1
Chrysobothris 5
mastersi 2 eo
Chrysomyia ae 325-6
albiceps .. 5 DA BAY
incisuralis = BAD, B40
megacephala 327-8
micropogon 327-8
putoria bo BAe
? rufifacies bo BAY
Chrysopasta 56 OZ
Chrysopogon so Owes
albopunctata > O94

Txexsxye

Chrysopogon crabroni-
formis 50 OS
fasciatus 50 OS
mulleri so OD
nigriscens so OD
pallidipennis BOOD
punctatus oo OS
queenslandi 50 OD
rubripennis 56 BOB
rufulus 5 OD
splendissimus st 395
Chrysorutilia ? 348, 352
Cimex lectularius XXVi
Cinnamomum ee Sala
Cisseis careniceps .. .. 228
SUB-BIFASCIALIS 29229
viridipurpurea ype
Cladochaeta 5
Cladolipes . 53
simplex .. so OS
Cladophlebis PLoS
Cladura 54
Claytonia australasica Veper Alar

Cleithrolepis granulatus lili
Clesthenia .. . 384
Clinopogon 395-6
Clydonodozus ao Oy!
angustifasciatus :. .. 54
Clytocosmus 58
skusei 58
Cnemoncosis ‘ 57
Cnidoglanis megastoma lili
Codium A 5 555-7
adhaerens 5 D5 Dae)
bursa Be) ga BOIS
difforme .. . 557, 559
GLOBOSUM 558
lineare so DDT
mammillosum DDS
ovale Sis cache Rone od ee Oe.
tomentosum .. x, bdl,959
Codula 2 sa oe
fenestrata ao oes
limbipennis .. 394
? quadricincta so aA!
? vespiformis So UE
Coelochondria .. . 56 DUK!)
Colobanthus subulatus so HAlis
Comesperma retusum
195, 204, 209
Comptosia OOD
Conosia 54
inronatal en. os:
Conospermum Shien PAS
Coprosma Nertera .. so Daley
Coptorhynchus ALBIVARIUS 354
jansoni .. 3b4
OCULARIS Soon:
TRIVITTATUS ODD
Cordyline stricta 56, alts
Correa Lawrenciana 56 US)
Cossonus 367, 376-7
Craspedia Richea ..
195, 198, 204, 209, Paley’
Richea var. alpina 217-8

1xxxvi.

Crocodilus porosus .. xlvii
Crouania . 461
Crypteria ; cow eh
Cryptochaetum 5 ela
curtipenne 50 OR
iceryae 422-3
LATIMANA a2 Oy
monophlebi 422- 3
Cryptolabis ao HO
Cryptopogon 5 BOG
Ctedonia a 50
Ctenacroscelis .. 58
Ctenocephalus' canis xxvi
felis xxvi
Ctenolimnophila. 53
bivena 53
decisa 53
Ctenophora 3 57
Culex annulirostris Ox
australis . 288
fatigans oe XV, XX, DGD-8
fergusoni . 565
flavifrons so CRS)
occidentalis oo CAS
procax ao cee
rubrithorax Ameer tE)CL
Culicada vandema .. .. 289
wilsoni 56 A)
Culicoides .. 2 Xe
Cyanonedys .. 394
Cycloderma ohiense 56) ADO
platyspora oe 256
Cyclophorus confluens ao BUR
serpens 5 DHS
Cyclopsidea 55 (Gee
Cylindrotoma .. 60-2
nigriventris .. 60
splendens so ED
Cynomyia .. 50 ‘axdil
Cyphogastra 50 BR
browni 55 Cs
farinosa 510 PR
macfarlani See ere 2,
pistor 5 LS
venerea .. Sa Les
vulnerata oad,
Cyttaromyia . 60-1
Dactylolabis ae aio FAY)
Dampiera 472-3 493, 495, 497
Brownii .. em Ashio
stricta 471, 473. sth 493-6
Danthonia robusta .. 5 PALA
Dapanoptera .. : 68
Dasylirion filifolius 272
Dasymallomyia 54
Dasymolophilus 56
Dasypogon 387
albopunctata 394
nitidus 397
princeps 397
rubrithorazr 394
salinator 397
spintha 394

INDEX.

Dasypogon suavis OOK
venno . 394
Dasythamnion . 460
Dasyurus viverrinus .. 550
Daviesia Cee ee 196, 199
ulicina .. 196,198
Dawsonia superba Se OLS
Deltastoma . 435, 442
unipunctata .. . 442
Deltopecten . 104
obliquatus : . 104
RIDNELS Tes oh las) see Os
Dendrobium . 570
Becklerii . 570
Mortii Later . 510
TENUISSIMUM a eae)
Depressa . 399, 400
ALBICOSTA . 401
atrata : 400-2
STRIATIPENNIS Sette hess Aa
Deromyia 395-7
Desmometopa m-atrum .. 8
VARIPALPIS .. 7
Dexopollenia chrysothrix 324
Dianella tasmanica .. 195
Diapontomyia .... .. 381
Diaspasis AN mee tattay eet | : S04
DiCKSOMLa eee en meee 10)
Dicranomyia so. oo (OES
_ diva aT CON VaR PR aed ta keke)
doddi 64, 67
THONG, 65 50 on WO
punctipennis Rey bere 14 OM
tarsalba Per reve batires YT OK
Dicranophragma i TAT AUM Ey OS
Dicranoptycha 63, 69
germana 68-9
Dicranota .. 48-9
bimaculatay |. sane ee
Dictenidia 57
Dictyosphaeria ‘favulosa BR 6
sericea . Pee eee Gey MON ae)
Didelphiy Smee ene o
Digenea simplex .. .. 555
Diotmephawe eae OS
Diplotoxa = Wee eho 4
microcera 434
TASMANIENSIS 434
versicolor é .. 4384
Dipodium punctatum 55 BUS
Discaria australis .. .. 197
Disciseda 235-6, 253- 4
anomala 239, 254-6
australis . 240, 254
candida .. 238, 254-6
cervina : 238, 254-6
circumscissa.. .. .. 256
debreciensis .. .. .. 256
hyalothrix 237, 254-6
hypogaea 240, 254-6
Muelleri Le 255-6
pedicellata . 2387, 254, 256
verrucosa au . 240, 254
DISCODOlA en oe ere nO

Discobolo australis ..

tesselata Per ike, (0
Dispar PPR NS Go BUD
Dizgonomyza é 426

Dolerothrips (?) GEIJERAE
57-8
Dolichopeza 59, 60
cinereas 22 |. See
mailagasva. > 2) Wasa
queenslandica a eae OS,

Doryphora sel an eee
161, 167-8, 172-6, 184
sassafras ; io abs
Drapetes tasmanica Apaear sky)

Drimys aromatica var.

pedunculata 192, 194,198

Drosera Arcturi bo Chilis}

peltata . 199, 204, 209
Drosophila Oates
albostriata

ampelophila .. Be 4

biarmipes iol eee 7

brunneipennis 4

buscki slo coes 3

BINIG@MA oc Sei Soe Ro

flavohirta as 3

fuscithorax 4) ee

hydei 3

immigrans 3

inornata 3 Ch tae 4

lativittata sig A regi eRe

melanogaster 4

mycetophaga ae 2

NICHOLS ONiIia a aner ee <4

3

6

3

3

3

3

6

3

6

5

Co bo

nigrovittata .. Hi

NITIDITHORAX nat oe ea as

obsoleta ..

poecilothorax

polypori

repleta PERSIA Fes:

SERRATA) 36 7.) aces

setifemur 6 he

SUBNITIDA Bol coteneecenede oe Cle

SYDNEYENSIS me Gs
Dryandra : 114, 258, 2
Dysmachus : a
Echidnophaga gallinacea xxv

myrmecobii XXVi
Egernia striolata xliii,

xliv, 25, 31, 499, 500, 553

whitei xliii, xliv, 25, 31,
499, 500, 520, 530, 553
Elaeocarpus ; 5 Wal4!
Elephantomyia 53, 63
supernumeraria .. .. 53
westwoodi ..:°°.. J. 538
ote ener mens. 0S)
OEP 56 6 i rey)
Elliptera 63, 69
AStISIMALICa) =e nO
CIAUSA ay ci Se OD.
illini weet Sao.) ie ee 9)

jacoti APG Rarattec ic. (UC)

Elliptera omissa the Daahi69
tennessa ore net A68-9
zipanguensis wa? Seb 9,

Elonichthys ALG

Empeda a orev ea Toes ae Ey OG

Empedomorpha atom okt)
empedoides .. .. .. 56

Enhypnon laticeps .. .. 231
TLONCHBATIGNE 565 ina oo ezeul

Eothrips BURSARIAE 5 Jus)

Kotipula .. Ree air

Epacris heteronema . 209
paludosa 194-5, 205-8
petrophila 215, 217-8
serpyllifolia 204, 206-8, 211-2

Ephydroscinis .. . 485, 437
australis Beene EAS

Epilobium confertifolium

216-7
glabellum 194-5, 197, 199, 212

Epiphragma setae oo: , OX")

Epitriptus fe BT dtotieee

Eremophila longifolia xl vii

Erethropogon 395-6

Eriocera 53, 55
fultonensis .. D2-3
gibbosa . 52-3
subrectangularis.. .. 53
velveta 52-3
wilsoni ~. Da-o

Erioptera 46, 54, 56
lutea ate So OO

Erythraea australis 197

IDOI, 955 oo oon Jia)
? BELLAMBIENSIS .. .. lll
BELMONTENSIS 110-1
coghlani xlvii, 105-8
forbesi 107-10
GLABRA a Id
GLENLEENSIS : 108-9
IPSVICIENSIS 107, 110-1
LATA cen re 6 Ld)
LENTICULARIS 5 LOY)
LINGUIFORMIS are pel ulbll
mangaliensis 105, 107-8
minuta A een O09
NOV OCASTRENSIS 109-10
OBLIQUA .. . 109
striata 5 alk
TRIGONELLARIS Be oo, LOY
WIANAMATTENSIS 108-9

EKuamphibolia 351-2

Eucalyptus 114, 161-2,

164-5, 180, 182-4
albens : Bi AS
amyegdalina var. nitida

184, 196
Blaxlandi 165,168, 172,
174, 179, 180, 182, 184
coriacea 114, 191, 194,
196-7, 200-1,
203, 212, 215, 219
goniocalyx 165, 167-8,

172, 174, 179, 180, 182, 184

INDEX.

Eucalyptus Gunni .. 192,
194, 196-7, 200, 203, 219
haemastoma 161, 173-4,
176, 179, 180, 184
haemastoma var. mi-
crantha oo. aUaxs Ural ley}
hemiphloia 5 CABS
meliodora MGA tS. eer}
piperita 161, 165, 168,
171-4, 176, 179,
180, 182, 184, 200

pre-coriacea .. 5 Jlite!
sideroxylon é Xxxix
stellulata .. 192,194,
196-7, 200, 203
viminalis 196-7, 203
Euchondria . 560
EUCOMPSOMYIA ae AD
LATIFRONS 325-6
SEMIMETALLICA 325-6
Eugenia australis 5 Or)
coolminiana .. .. .. 379
Euglochina SRN bee ey OF!
Euhippelates 435-6
pallidiseta 30. 40
Euhomoneura . 419
ornatipennis . 419

EKuphrasia Brownii 5
195, 204-5, 217- 8

Euphumosia 5 KDR, 3783
calliphoroides .. 324
eristaloides . 324
papouana .. 324
papua 323-4
variegata : aN . 324

Hurhamphidia .. .. .. 69
niveitarsis .. 58-9

Hurina 2» 442

Eusargassum .. .. .. 560

HUSTACOMYIA ..) 45.25 337,
BREVISETA Fy 1 OREO RT ERR OE

EUTHYRIDULA oo etl
rugosa is 444.

EKutolmus D0) ail

Eixeretoneura 5 Gxsil

Fagus .. MEAS heya weles:

Fannia canicularis wie XXIV

Ficus eugenioides 5. Os)
henneana Botait oe be axthee)

Fieldia 5 AL AG
australis . 174

Filaria bancrofti XGV I EXLOXS

Finlaya an . 298
alboannulatus eb 65
notoscriptus uke 565
queenslandis AG 298, 565

INOS, oso 56 oa. oo GOA)
atribasis Sybil lucies 350-1
flavipennis .. .. °..+ 350
mirabilis i Dae 350
speciosa .. ; 349, 351

IDOVASAANOUIE, Go 5a oo oo Zhe!

Hranklandiay) {sens ae 2D 8

Ixxxvyii.

Galium umbrosum ..

194-5, 197-8, PAT
Gastrodia sesamoides .. 198
Gaurax . 4385, 437, 444
APICIPUNCTATA . 444
atrisetaiiye, Sera loos sea se
Geaster , ‘ 253-4
affinis a OCA Ae RO
VAN CIICT Aeneas. eS) Tee OSG
arenarius A rare eae
argenteus cae SRN eo 6
asper aed Sano
australis Set eee eee
Berkeleyi Lay BAS Sch eee FA
biplicatus Sevens 256
Bryantii : 254, 256
CACSDULOSUSH ee ee 6
COLCEUSOAR, 05 eee a6
campester 255-6
Clelandii chy, oe ee
coriaceus ives See ED O
coronatus dit) ZG
delicatus 50 CAMO
Drummondii 255-6
dubius bend NOR en 27215 6
ellipticis SU eh tht aaeerrqa ie
LH OPA MOOUS So 60 oo PALS
fenestriatus 5 255-6
fimbriatus 5 CA ADT
floriformis 255-6
VOTMMCTUUIS a5 66 ao AXE
granulosus 6d.) ace CAND
Hariotii Sa) Wane yes Bee
hungaricus 56 og 7D
hygrometricus .. ... 256
involutus iE) lie Su bee Gs
lageniformis ein 22 16
lignicola at Psa ye ola MerAS Me)
limbatus Bete st aie 15)
IGUOVOHE. ac 90. 40. 06 ZG
MAMMOSUS ey ee
MOAERLOULOLLS ae) eee O
MinimuUs iS, eae e256
minus 255- 6
mirabilis 255-6
Morganii » 256
orientalis ee cat ioe aD
pectinatus 254, 256-7
plicatus .. . 254, 256
pseudomammosus e256
Readeri BG) roeL CAO
rufescens neous oar c)
_saccatus Paes barat 700)
Schimideviiw a. sea) oO
simulans Se preteen te DD.
Smithiis a) cee eaneiee DD
striatulus Bede eon aes
SHACTUST Roe
subiculosus .. .. .. 255
tenwipes 256
triplex "255-7 7
tunicatus Me Wits d ot eure Ty
ummobilicatus .. .. «. 2d
velutinus

1xxxviii.

Geaster violaceus so ZB
Geijera parviflora .. 157, 159
Geoplana : XXXVii
Geranium dissectum 195, 197-8
Geranomyia Ry ears TON.
picta 66-7
tonnoiri .. 66-7
NGI bo sae og foo, We
victoriae 66-7
@itomidesiees n\n) ecco 7
CONVERGENS .. rk

Glaphyropyga australasiae 392

Globularia .. 484
Glossopteris 109-11
Glycine clandestina eg9
Gnaphalium purpureum 379
Gnophomyia Poti Vena Buses CPs):
Gonomyiae oh oe 2514-60
Gonops geniculatus 390
Goodenia .. .. .. 472,493
cycloptera ne ehO'5
hederacea 198
Goodia lotifolia 379
Gracilaria taenioides 555
Grapholostylum 352
decorum 48. 352
dorsomaculatum eh eas Die
Grevillea X6:O0b:¢; xd, 114,

122-3, 125, 258, 260
SCAM MMOIE, oo 06, a0
122, 124-6, 128-32
acanthifolia var. querci-

foliae) ss. 5- Veo at teal!
australis 195, 215-7, 259
buxifolia ne 125-6
Gaudichaudii 122-5, 129-32
lanigera 2) See 6
laurifolia 122, 124- 6, 128-32
parviflora Bo AAG
phylicoides . 126
punicea .. ; 126
sericea 125-6
sphacelata . 126
Griphoneura . 429
Gryllotalpa . 455
Guioa semiglauca no Ue
Gymnastes 51, 54-5, 69
ornatipennis.. .. 55
Shirase yee OD
Gymnobaris pA RUM 751:
Gynoplistia <3. gee O
bella ia 2 aie VOR ALIEA DO
dimidiata 3.8, OA ee DL
RIAU TAS Terese g se oa
hiemalis ieee | Waren Rab
HASvOMItIC’, 2. “ac ieesol
EVADLOmMastix, 42.6, neous
Habronema megastoma xxi
microstoma REX
muscae 5 OS!
Haematobia exigua AA >.©.4
Hakea 5 A425 8, 260
ambigua PN (fe

INDEX.

Hakea amplexicaulis . 269
arborescens : 50 AAD}
auriculata . 260, 269
Baxteri . 265
Brownii . 264
Candolleana .. .. .. 266
ceratophylla . 260, 265
clavata . 268
conchifolia Sn enone
corymbosa 5 ADIL, AZ IL
cristata .. . 269
cucullata 260-1, 272
cyclocarpa : » AIL, AOS
dactyloides 5 ILS, ABO, AU
epiglottis . 259
eriantha ye 20
eucalyptoides a orAtal
falcata 5 ADT
ferruginea . 269
flabellifolia See 04:
florida 5 CADE
glabella .. . 269
Hookeriana Beet ee ZO
incrassata . 260, 264
lasiantha 55 ALD
laurina 271-2
linearis . 269
lissocarpa BE Big cA)
Macreana 195, 198-9
macrocarpa . 259, 263
marginata 268-9
megalosperma .. 268
microcarpa 199, 211
Morrisonianum .. 267
multilineata .. A 262, 270
neurophylla 5 MA
nitida aA ee OTS
obtusa . 262, 268
oleifolia . 5 A
orthorrhyncha . 259
pandanicarpa 267-8
petiolaris 271-2
plurinervia a BUA
pugioniformis . 259
pycnoneura . 267
Roeii .. 264
ruscifolia 269-70
saligna . 270
stenocarpa 3) AUC
trifurcata MMA LED gh PAS)
undulata 2 2603 20Al
varia : 6 ZAG
varia var. intermedia 273
Victoriae aa AUC
vittata . 259

Halimeda > DDT
incrassata > jaz
ODUNTIAN en) se 5 atl
POlYAACEYLIS! |) eID DI
tuna | 555, 557

Haliseris , . 560
acrostichoides . 560
Muelleri.. . 560

Harrisomyia 53

jo Ut

Hebenstreitia
dentata .. Saini
Helichrysum baccharoides
194-5
bracteatum Ro di. 1}
rosmarinifolium .. . 194
scorpioides 195, 197-8
semipapposum 197-8
ee aa Ae 5 aril
lascus : 5 OY
Helipterum incanum 217-8
Helius us 63, 69
uniformis i | Ee OS
venustus 68-9
Helix aspersa .. . xlv
Helobia Re os ae) 8
Helosis 484

Hemarthria compressa wie
204-5, 215, 217

Hemicistela testacea » 2e2
Hemipyrellia 55 OrAl)
cyaneomarginata . 320
fergusoni 5 Al)
ELE NDELONSY.ZAU) 4) ene 8
HU DOMCOOMNHIS Go 40 oc 8
Hesperilla .. 275-6
andersoni 2s
chrysotricha . 279-80
chrysotricha chryso-
tricha . 280

chrysotricha LEUCOSPILA 280
chrysotricha PLEBEIA .. 281
erypsargyra .. sil:
crypsargyra crypsargyra 281

erypsargyra HOPSONI .. 282
eyclospila 278-81
donnysa.. 278-80
donnysa AURANTIA 5 UY)
donnysa donnysa o ang)
donnysa FLAVESCENS .. 279
donnysa GALENA .. . 280
idothea » BUS
picta 281-2
Heterangaeus 48, 50
gloriosus Semis Liter: cts
Heterolimnophila .. .. 50
Hexatomia: ise 2) eS
megacera syayl.) Vet eS
Hibbertia linearis var.
obtusifolia 195, 197-8

Hinulia quoyi xliii, xliv,
25, 30-1, 499, 500, 553

Hippelates .. 485, 437
ABBREVIATA . 437, 440
ATRICORNIS 437-8
atriseta .. 437-8
bancrofti .. 437
FERGUSONI 437-8
NIGRIDORSATA 437, 439-40
UNIMACULATA 437, 439-40

Hippobosea camelina . XXV
equina 5 LOY

Histiopteris Ai trees alte
incisa . 162,181

Histriobdella homari liv
Holorusiays.) .2 ant LOS
Homoneura 12, 400, 419
- apicinebula 13, 420
armata . 13-5
ASYMMETRICA . 13-5
atrogrisea 12, 419
barnardi iB}; 1s)
eidsvoldensis Biehl tees OL ka}
FERGUSONI .. 420
FLAVOFEMORATA . 420
fumifrons 5 Rear ate eeaeaa a Ls)
GORDONI .. 13-4
ILLINGWORTHI .. 13-5
indecisa se lord
ornatipennis . 419
PERTHENSIS . 13-5
preapicalis 13, 15
proximellay 82 se es
signatifrons .. .. .. 13
TESTACEA ‘ . 419
Hoplanthemistus albo-
fasciatus oo. 7%!
conifer . 234
Honistomyila 2.25 '.. .. 50
leucophaea .. . 56g DOS
Hormosira articulata . 560
Hornea : : 5 Sabb
Hovea longifolia 5 NG, sus)
Hudsonia .. peel nl PGs)
Hybrenia angustata 5 RY:
ILLIDGEI .. >, oe
pilosa j . 234
Hydreoclathrus cancellatus 555
Hyla cerulaea .. XXXVii
ewingi F XXXVI1i
Hymenanthera dentata
198, 217
Hypericum japonicum 198-9

. 555
. 229

Hypnea seticulosa ..
Hypocisseis suturalis
Hypolaena lateriflora ..

199, 203-5, 209, 212

Idioglochina .. .. .. 67
TaKoryowe, 6 be “oo oe DY)
Idotasia ae 5 OU)
llex 5 > Atal
Incurviseta .. 400, 402
AFFINIS .. 402, 404
BISERIATA .. 403, 406
FLAVICEPS .. 402, 405
FLAVIPALPIS .. 403, 407
FULVOVIRIDIS .. . 403, 407
LATIFRONS : 403-5
maculifrons 402-4, 406
TASMANIENSIS . 403, 406
VIRIDANA 403, 407-8
VIRIDULA . 403, 408
VITTIGERA . 403, 405
WILMOTI . 402, 404
Ipsichora so a)
BIMACULIBASIS so Oho)
desiderabilis 369-70

INDEX.

Ipsichora TIBIALIS

Irona ..

Ischnothrix
australasiae ..
connexa..

Ischnotoma

Iso flos-maris
rhothophilus

Isopogon

Isotrogus bilineatus

castelnaui
TRREGULARIS

Jania rubens
Juncus pallidus
pusillus ..

Kladothrips
AUGONSAXXOS
rugosus ..

Kreyssigia multifior a

Kunzea Muelleri

57369
xlviii
52-3, 63
52

52
DS
5, dbrbs<
xIviii
P2518
5.6 BUT
oo BUT
5 UO

.. 555
co UY)
> IY

153
“153-4
153-4
. 378

195, 199, 216-7

Lacerta agilis .. 503-4, 506
Laosa .. Mae OS
Laphria . 094, 396
Lasiothalia 460-1
hirsuta A 460-1
Laurencia obtusa 555
rigida 555
Leaia .. 105
Lechria 62
singularis 62
sublaevis 62
Lecteria . 54-5
Leiomyza 445
LEPIDOBARIS 370
METASTERNALIS 371
Leptoconops xxi
Leptogaster 389
aestiva 390
antipodus 390
australis 390
autumnalis 390
bancrofti 390
dissimilis 390
fumipennis 390
occidentalis 390
pedanus .. 390
vernalis .. ‘ 390
Leptoplana australis liii
Leptopsylla musculi XXvi
Leptosiphon 484
Leptospermum ppel99)
lanigerum 194-5, 198-9
stellatum 196, 198-9
Leptotarsus Bas SUR'S
macquarti ao BN
Leschenaultia 42
Lestophonus sig AAIl
Leucadendron 8258
Leucophenga 2

1xxxix.

Leucophenga MINUTA .. 2
Leucopis ave 5 APA
Leveillea jungermanni-
oides OD
Libnotes 64, 66
STSCI 4 se Se 66
OQOWCHA. 65> oo sy on OB
TC LATS CY, cone ade ee OS
ROSIN grein wwe. EE LOG
subaequalis .. .. .. 66
thwaitesiana Cuneeetab6
undulata a ae. Borie ad)
WiCiN OM aa eS
aerea NOS TE reer ae iT
angusticollis A MeO
apasioides ys al eo
Coeruleaicy yc 5) ee
commoda 231
cyclocollis 231
elata 231
gilesi 231
illidgei 231
major 231
meridiana 231
monticola 231
nitida 230
nitidissima 231
nodulosa 230
pallipes .. 230
planata 231
puncticeps 231
splendens 231
subcanaliculata 230
sylvicola STE 5 eb rad Be
tasmanica 230
truncata 231
umbilicata : ag 2D)
Limax queenslandicus se wall
Limnophila 50, 53, 62
Limnophilella ... 50
imoniays we ee. | ee
amabilis Ler IRE Al ae oe tl
PINAL TK eee Me ete en eee Os!
BOOTS) 64 oo oo oo BO
Eeenclorsibon 55° 4 so (87
basispina Sek Sea OM
bifasciata ite ae SPO
CalifornmiCa ae nO
cinctipes Yeu Na es 67
esakii cS scat Ne SRSA eee ok ONG
fallax Ar Mums me ts) Li'sch bY Natl
HAV IDES celeste eee Ord
fumosa 66-7
immatura Liay > jaan eae Ont)
indigena SEAN Sea aise PONE!
insularis ee Sete ot tr Bay
LIVUS CAM ae ay, ere Seen ON)
TMEKCWINKGOSNY o5. ac oc al
THOT OOWUORE, 855 oo 60 WY
parietina Ae SLE Oe
RPA, Go oo bo ho
quadrinotata Stan Git
sciophila SP ectal aa Ot
solitaria Sete tay ilk. fai

xe.
Limonia sponsa Se RS NONE:
triocellata PUNY 2 Lo escn 3 PLS
(BEYONCE, sa oo oo, Ol
tristigma NAN ms ob ae Peer’ OV
tristigmata 66-7
trivittata ORE dees SUL
Lindsaya linearis .. .. 81
MMNGRO OAV, co oo bo til
Linum Be uebeTEs cs ets oaks ore
marginale 194-5, 198
Liogma Ales . 60-1
glabrata 2 Rom es ee Gill
nodicornis Lat Lol etait geet batt ()

Liolepisma entrecasteauxi,
xliii, 25-6, 499, 500, 546, 553

Liphyra brassolis .. .. liii
Lipophleps ees eb leetay Sh 15)5)
Ib OW ME oo) Gea doo sau, Be
Liriomyza oper He Ny42 6
PALLIDICENTRALIS .. 427
OMG, 56° “Bev e0 oa Aa
TRICOLOR eee sk CLA
Lissanthe montana 195, 216-7
Lobelia . «. 484,495
trigonocaulis Oe Be. Be
Lomatia 4 alas, eal
longifolia 196, 198-9
Longuriv .. AS
Loranthus pendulus 50. BUS)
Lotus australis 197-9, 204
corniculatus ~ LOT; 199
Lucilia 303, 316, 318, 320-1
caesar i 321-2
GRO oo oo da loo Sul
cyaneocincta fDi:
cyaneomarginata .. 320-1
fergusoni Bre ene Vig os OAL)
FLAVICORNIS 321-2
nosocomiorum hee 321-2
pallifrons gate Pa Seat eee AL
SCLICALA Coe
Lutzia halifaxi FER ee dt COMED)
Luzula campestris .. 204, 212
Lycoperdon oe. AEDS
asperum 254-5, 257
QUST Alera e 257
BOUUSTO acs 2 EL Ae
POVAIStOIdes!);, ee oe
candidum PM Ll ns hrAD
CEPACTOTFANE .-)) > eee
Colensoi Mo Oe foc PANT
coloratum i 11 BS.
COmpActim, =) | See ec DS
COOKE. 50) eee
coprophnilum.. Peas
COLIN os MERIT
cruciatum day vee aD
depressum .. .. 253; 257
dermozanthum ..... 257
fUUOSIIVE 2) et eee OLN
Fontanesti. .. ..° .. 257
furfuraceum Pee by hilt
gemmatwum .. 2.215.. 257
PiPanteum (7). | eae. edt

ENDEX.

Lycoperdon glabrescens 245-5

Gunnii 254-5
WOCHUBUG o6 566° on. 06 ABU
hungaricum .. .. .. 257
lilacinum Se ccatevemieres e 2 Dyl)
macrogemmatum so ZANT
microspermum .. .. 257
natalense eA aes TG
OCU 06 o0 ‘on oo Gadd
mvenGlbeat G5 6k oo) CU
novae-zelandiae .. .. 257
perlatum 5 253,257
piriforme 5 ADB, Ze
piriforme var. flavum 257
polymorphum . 2538, 257
pratense id chen PEAR e ON
pseudopusillum .. .. 257
purpuraceum ae cee
(DOW OURGOHE oo a5. oo ADY
pusillum . 258, 255, 257
TFOLLCOMETHT 56 900 60, HDT
retis A sf Leet a BT:
rubro- flavum | Riera 45) |
scabrum Bie 254-5
semi-immersum .. .. 257
Sinclairii Dele Pleas eke Sl
spadiceum .. .. 253, 257
stellatum a NE PODS
subincarnatum 5 A, CADE
substellatum at 257
tasmanicum ROR ao ea
tephrum Prarie poss? ca
violascens BUSTA NEMO ay LeU
Wrightii ae 2 YG
Lycopodium clavatum on ALT
Selago .... PAT
Lygosoma entrecasteauxi
xi, xliii, xliv, 25-30,
499, 500-1, 503-7, 525-6,
543, 546-53
quoyi xi, xliii,

25, 27-31, 499-501,
503-7, 525-6, 543, 546-53

IMachimusiy eee SO
Macrochile 5 Soe a plete he Nee Ally
Macromastix. 2 phe sae DS
COStALIS He...) cna BS See eS
Macrotaeniopteris .. .. 108
Mandalotus Bie cron: eae Oe)
ARMICOXIS ice ppeney breeehere iT
campylocnemis .. .. 356
CaLnInatipesi i aE So
HourseoreNbI 55 55 oo BHT
mirabilis oo RE) Geo
Dyer >) See obo
STERNOCERUS SRE ay or CLG
Viet VEUIS) heer), |) Meio OO:
Maria bide as Seto 4
Megistocera ee sadrratn axe)
longipennis 58-9
MeESISTOMAStIX. arlene OO
Melanagromyza yess esa

ALBISQUAMA

Melanagromyza phaseoli 424-5

-TRISPINA 424-5
MELANINA .. - 400, 412
AENESCENS! (5.0 cele
MAJOR 412-3
PLEBEIA 412-3
QUADRISELIATA 412-3
Melanterius oe ee OA:
Melinda Ae bow hOB
Melophagus oy inus BEPXOXGY,
Menyanthes Peat, prod aeKe
Merimna atrata Rie Sic ee eae
Meromyza Ms oo, CIS
Mesodina .. 22) Ba Aae275
Metalaphria é 395- 6
aurifacies Li Saas 96
australis a Ce OOL6
Metallea 2. | :)) sah o e829
gracilipalpis .. 329-30
illingworthi . 329-30
INSULARIS 330-1
nigribarba 330-1
PUNCTICEPS 330-1
robusta 329-30
Microcalliphora 325-6
annulipes 2150s hae nem a O
flavifrons eee so AD
VOUOYNES 56 60 00 oo S28
Vari pes! :c..We eee 6
Microchrysa io oO
Microtipula say) eee eee
Microtis porrifolia .. .. 378
Microtragus 6. DE Oe.
arachneias se Oe
arachne var. sticticus 234
CEUSTUWLOLLS tS:
echinatus aia t he 234
Microtropeza "'952- 3
Miltogramma:.. .. .. 335
Mischoderus .. . pee er! 45)
Molophilus “46, 54, 56
froggatti Brod Diemer tim 83
Monophlebus.. .. 423, 446
Monospora ee the eae O)
Mormoniella brevicornis xxiv
Morphomyia .. .. .. 339
tachinoides .. .. .. 340
Motasingha shes) Levey Rape ie AHO
dominula at) Src:
drachmophora Pree. atts:
monticola shai. | Saicet nee
Mugil ae . xivi
Musca nate - eo Oe
domestica xv, xxii, 302, 314
Mycenastrum 245, 253-4
bovistoides) Ys) eee
chilense cele rosie MR we OI
clausum Rr siar
corium 245, 254-5, 257
leptodermeum a 257
olivaceum 1/5 Soiree Ot
phaeotrichum erie Ane
radicatum Sint Late) Ciena ON
SPUUULOSTTI weld ethene ti

Mycodrosophila Siesl (ie 1

ARGENTIFRONS i if
Myctides balaninirostris 366
Myoporum .. 484

laetum xl viii
Myosotis australis Pei Scicer tly
Myrica ies age ele:
Myzomyia .. oo-4

amictus .. 34-5, 37

annulipes 5 OOD)

mastersi chit MR haa O10)

punctulatus . 33-4
Nasonia brevicornis xxvii
Neoaratus 385, 390, 392-3, 397

inglorius : 393
Neocalliphora j 304, 308
Neocerdistus 390-2
Neocladura ae eite ent Bye
Neoculex fergusoni so BOD
Neocyrtopogon 383, 395-7
Neoderus .. .. . 45

patagonicus .... .. 45
Neodioctria 396
Neohesperiila .. .. .. 275
Neoitamus Ad an Bol, Bol
Neolimnobia .. .. .. 68
Neolimnopfiila Se atpan aye
Neomeris dumetosa 56 BOO
Neopollenia . 3804
Neosarapogon 395-8

Claripennis) 2. 3. .. 39

froggatti Bae HS elitacas oneself

nigrinus fey Meh Aerts teen ts 91:

princeps 397

Neospades chrysopysia 56 eve)

CUPRICAUDA .. 228
eOuGH: 65 bod oe ae. 24s
lateralis te 228-9
NIVEO-SPARSA ae oo | AS
Nephrops norvegicus eo Ibhy
Nephrotoma dts 58, 60
INGEOWOZE 46.) bo a0 oo OY)
Nipponomyia 47, 49
kuwanai nbc abe ah eT a)
novem-punctata .. .. 50
SOMATA, | “go ao oo ON)
symphyletes .. 49, 50
trispinosa Ree eee AP) ()
Notaden bennettii .. xxxviii
Nothoderus Se rece iat ene) 92:5)
australiensis fe rey
Notiosomus 5 Se
INSULARIS eM OO
Nusa : . 394
Nyctozoilus pemionaivencis 232
TAYLOR | 32 1 ao CB
Ochlerotatus . 284, 286, 288
aculeatus . 295
albirostris . 288
andersoni ~ 291
antipodeus .. .. .. 288
burpengaryensis .. . 294

E

INDEX.

Ochlerotatus camptor-
SARCINUS 56) 50 on Zab
clelandi .. yee tine 294
cunabulanus.. .. .. 292
flavifrons .. 289, 565
imMpRimens Meme eos
luteifemur ney ae IO
MACLEAYANUS At oe Oe yl
nigrithorax Pete eR age
moans oo. oo do oo ZAYB
normanensis 50. ZNO
purpureiventris -. 294
rubrithorax ao Ae
sagax j be 292
stricklandi oo XD)
theobaldi 5 oo OD

theobaldi form &EIDS-
VWORDIWSHS oo oo oo AD
vigilax . 296, 565
vittiger oo Pe)
Ochromeigenia 5 60 OO
@immONONGleS 46 oo 9 oo Bests
Ochromyia 56 0 6 ON
2? hyalipennis oo. oo alts
Oenothera biennis .... lili
OCOPANE, oo ba 06 | oo lb
Oestrophasia Pay Ese: Lave al)
Ogyris aenone.. .. .. 1
amaryllis OUEST E LT Ae 1
OTOeCTES We le ue fe 1

Olearia flor bunda PAILT(
megalophilla 194-5, 198, 200

myrsinoides .. 192.195, 198
Ommatius as x 390
Onchocerca eibsoni- XX, XOX

gutterosa Hig Sine aan? a

lienalis 66 Oc
Onesia 301- 2,316
Ophiodesma a Leo sate &
Ophiomyia . 424, 426

lantanae . 426
Opseostlengis .. .. .. 395

insignis She (bau. LORD
Oreisplanus 275-6

Oreomyrrhis andicola 195,
198-9. 203, 206, 212, 217-8

pulvinifica 5. Ake
ONCE, 55 of oo oo We
alpinay tases. ar ee O64
IGN Ae es, ere eee ats OE
damped i G4
excessiva 5 63-4
fORIMOSICOLA NEE nO
joana a 63-4
TAIAWEMEBIESNIS 55 6a) WE
TOGWURIGINIORISHIS Fo oo 06 (eh
punctipennis 63-4
Orimargula 63, 68
australiensis 5 56 ASG)
Orites lancifolia 195, 216-7
(ORMNORIEY 55 “co c co DO
Ornithodes ca. Wnt Poet
Oropeza NS ALAS OO
Ovinus mylittae 55 DUS

KCI

Oxylohium ellipticum var.
alpinum 192, 194-8, 200, 208

(HOUIOORNADION oe o6 oo ord)
OVAOCKIOEIE, 556 co co da bis
Pachylophus . 428

ALIENUS .. 428-9

lutea 428-9

SECUNDUS 428-9
IEAGUTENOINE) a6.) oo oo os CD
Palpostoma 337-8

APICALIS Silat yal Lee oo,

desvoidyi OOO

testacea .. 56, asaxe:

Panax sambucifolius 194- 6, 198

Papilio aegeus .. xlvii
aegeus ormenus xlvii
androgeus xlvii
oberon . ao xlvii

Paracalliphora ero

Paragymnastes 51, 54-5
haASCIpPeninilSee see DD

Parahippelates - 435, 437

Paralauxania 400, 408-9
ATRIMANA 409-11
elevata 409, 411-2
FLAVIPALPIS . 410, 412
FLAVIPENNIS 3 409-11
fulviceps . 409, 411
NIGRIMANA ab 410-2
SCRIPTA ae 409, 411-2

Paralimnophila Besa pares Kona!)

Paramenia : 5 aby

Paramphibolay s..5) so len ene

Paranomina . 400, 402 -
unicolor aot ae Gh

RaTraracusmeacwaee . 390, 393
MACLOStYAUS = VoLe Ooo

Paratricyclea .. .. 303,322
TaORKEhieWagees Meee pts.) th LOR
SUIRGOWNT 55 oo oo, oo RS

Paritamus Lat Chere

Pectinotipulina Rhy Mest taarss

Pedicia ; EA 49, 50

Pelecorr hynchus EO OL!

Pentachondra pumila .. 217

Pentarthrum : 5 BU
australe PA ols, Sao
FOVEICEPS aM 374-5
FOVELVENTRE 4. 9.2 2 ole
huttoni ne 372-3
INTEROCULARE eo OD
MiUillimMetOnie MOS
nepeanianum 5 Ba
nigrum 373-5
ORTHODOXUM .. 373-4
SMPMCIKCOIEIU oo co oo OD
zealandicum .. : 372-4

Penthoptera TER OES) eae E183

Perameles ..

28-30, 499, bad, 546, 550- 3

Peringueyomyina .. .. 45
barnardi RA TEE EA 5

Peripatus XXXVii

xcil.

Peripheroptera 64, 67
CUCORPO® oo o6 60 05-2 WY
HOVEOUNINOG, so oo oo Wd
TOSIOSY | So gs Seige ton | oo oO

Persoonia re ee onset) La yt
ROO) od! be a6 oo Jet
chamaepeuce 197-8
Fuanyoeenine, 66 bo oo Zo)
INM@BTENIS og. oo a0 oo BUY)
salicina 176-7

Petaloxylon porosum .. 113
scalariforme URLS

Petauroides volans xlviii

Petrophila Te ae ey TaD OS
linearis .. . st EO An)

Pezohelaeus denticollis . 5 OW)
WnEUS Aeneas Beet)

Phacelodocera BUSA LEDS

Phalacrocerd eee le nOO
formosae Pata Bare toO
OCCICENntaliShey ce COO
replicata . 60-1
tipulina . 60-1

Phalacrocorax carbo tees BL

Phebalium ovatifolium ..

194-5, 216-7

Phellus . 388, 394
FIBMCUS 56°00 ob . 65 Soe
piliferus SA aioe.

Phenicia j 320-1
CUI, oo ca ao 40, Call
Sericatan ey. Bian te eo2r

IPnerocerawae i | eos

Philonicus 385, 391

Philydrum lanuginosum 378

Phlaeophagosoma .. .. 376
corvinum fee, oA Dee OTG
chulrpAvbea, 26° 56 on. oo. UD
pedatum ee OT

IAN OKO = 55 be oo 2x

Phumosia .. 2 a2Z0 3823
ANAS eee eee Oe}

Phyllolabis 50, 53

Phyllotheca Fiche cas) eLOS
australis OM LE cee Ope)

Phyllotricha He Niaak eigk One)

Ey Mmatopsisi. mee ees

Pimelea alpina 216-7
axiflora 5 ARYA TUS a, ale!
axiflora var. alpina .. 217
SLATI CA es a coand, eae MELO.
ligustrina 366 Bley aaa 2
ligustrina var. hyperi-

cina j 195, 198

Piophila contecta 8
latipes ...2 tt A 8

Pittosporum rev olutum ong

Plantago varia 195, 197-8, 212

Elatycephalus... Assit. oli

Platycerium grande Se Loue

PLATYINA ; 435-6
WEBULIBER AM Wee Us 1436

Platyiimnopia.. sail aie

Platyphasia sete ob Me ABS

INDEX.

Pleonosporium ey re 4 Ob)
brounianum .. . 464

Plusiomyia Ata eeaeeminis in: | LNs:
felix eee ber oies (LL ede 8

Poa caespitosa .. .. 192,

195, 197-8, 204-6,
209, 212, 215, 217-8
Podocarpus alpina 195, 215-7
Podolepis longipedata ..
195, 198, 218

Podoneura Se TG
Poecilohetaerella . 400, 408
Poecilohetaerus . 400, 408
decora Ne . 408
punctifacies 66. oon COS
schineri aba OO RERAYS
Poecilopachys bispinosa liv
Pollenia 303, 318-9, 328
auronotata .. .. 303
HIRTICEPS Sieh larat sd hhs VESTS
moretonensis .. .. 335
rudis ee Mee els waa aa eee As [tet
ruficornis Se) ee Ero D
tasmanensis.. .. .. 335
viridiventris aS 335
Polyangaeus “48, 50
maculatus J RB es Teg
IPO ANE. 66 - oo “oo. oo 0)
inornata arse > Uae Meena sty ()
Polymoria red eect) ()
‘Polyporus mylittae ETS
Polystichum aculeatum ..
195, 215, 217
Pomaderris Disk eae ee thes LL
Poranthera microphyila
195, 198
Prasophyllum fuscum-.. 203
Prionocera Sch aeewe | eth OS
IETPIOMOUB, oo. 6a oo ac Be
Prodiaphania .. .. .. 351
testacea PHA tal Gin)
Promachus 390-1
laciniosus wc (eat eer Oli
PROMUSCa Sh yo eee Oe
Prostanthera cuneata ..
194-5, 216-7
Proteosoma 7 MONEY ek PES OKAVE
Protipuiay s.:, '/2c, £590 eee
Protomiltogramma ooo
Protoplasa sie Sehyeish ab ae
Prunella yulgaris 197-8
Psaronius .. 51, 55, 69
abnormis oe 51
brevitibia ee Weaee? Seta:
MATTCUS aes ts ee IL:
ODIITEKATIS |e eae
obscurus Par bA act ede, Aun etiay Ul
trianguliferus nM alas etal
Pselliophora Sa? 57
Pseudolimnophila 50) 53
Pseudopalpostoma .. .. 339
Pseudophryne australis xxxvii
joniorwonahin 45% ; XXXVii
Pseudoskusea concolor 5.0 PAD

Psiloconopa Renae Hob okt
Psilostoma incisuralis Bi OC
Psilozona .. 232 Ove
Pteridium .. 165, 167, 172, 184
Pterochionea .. .. 54
Pterohelaeus .. .. .. 230
WOHMOHHIS 556 60 co on 2A)
sueriniy s. | aye oO)
puer Sk eee ZOO
CU POGUHTOS oo oc oo atl)
ventralis id Eee 23 0
Pterostylis acuminata .. 198
coccinea BTM. 43) Le
mutica sic Gs See
Pterycota.. i. oo eee
Ptilogyna 58, 60
Ptiloniesiay 3.) 2) eee Oe
aunonotata 25 eels
Pinlostena 4-0) sae eS
Ptychoptera lenis .. .. 70
Pulex irritans .. XXvVi
Pygiopsylla rainbowi xxvi
Quercus a a aaa alt
Questopogon 395-7
QOuintinia ~. 3. fa ee 20
Sieberi 3: 3. aS
Rachiopogon 395-7
Radinoderus 54 hen
Ranunculus anemoneus PALL
dissectifolius che PeeereaevetlaT.
Gunnianus 217-8
lappaceus i) Se Lg
Millani MS hire aca “dle
Muelleri nie 217-8
rivularis . 216, 218
Raoulia catipes 216-8
Rattus norvegicus .. .. XXV
rattus . XXV

Restio australis ae
203, 206-7, 209, 212, 218
56

Rhabdomastix ..
Rhabdotoitamus el ROL
Rhacopteris aoe 55 HRA
Rhagadolyra .. .. 400, 413
handlirschi .. .. .. 413
Rhampholimnobia .. .. 69
Rhamphophila ha ee O
Rhaphidolabis .. 46-7, 49
CAYUGA ch lie | a
major er caro CL
polymeroides <a SAS
rubescens Pre 3.1, ANE
stigma 25.. 25 ee eee
Rhinia ap a oe BAS), Ba
quadrinotata : Se io,
Rhinoscapha INTERRUPTA 355
CHICOLOT ty ee) ere O
Rhipidias ci. eee eONG
Rhynchodemus XXXVii
Rhyncomyia .. .. .. 329
dubia 329, dad
INCISULALISMe se Eero

Rhyncomyia tigrina 50 MOND
Rhynia ato > pdllibe
Richea Gunnii.. .

194-5, 204, 207-8, 215, 217-8
Roestelia polita xvii
Rubus parvifolius .. 196,198

rosifolius Bes, Sette seei9 5:
Rumex acetosella ..
194-5, 197- 8, 212

Rutilia 342, 345, 348, 350, 352
aneentiterd 5. <<. .. 349
GeCOnameesy see ise. Pe sab2
desvoidyi 345-6, 349
erichsoni 348-9
formosa .. 347-9
inornata . 847, 349
splendida 347-9, 352
vivipara .. 345-6, 349

Rutilodexia Pace ee lame aay.
angustipennis po. 6g De

Sambucus xanthocarpa .. 378

Sanmvalwmi ss a se eld
Sapromyza ,
8, 400, Agile 3, 415, 418
AVICOLA , 416
barnardi mes Pelanee skh, Phy
BREVICORNIS .. .. .. 10
brunneovittata .. .. 12
flavimana ae rag kee Seoretiy PLL!
FLAVODORSALIS Sena ewe alle)
HIEROGLYPHICA ..... 415
maculithorax LN eee Dy
magnicornis .. .. .. 9
magnifica Beth tak, hiccty a weil)
IMUNRDAI Ve cel wos ee ll
occipitalis adeel FAP ammo te U7
OCELLARIS Oe estan kali
PETERSENI 414-6
PLUMISETA 414-6
regalis 416-7
RIPARIA .. 9-10
sciomyzina .. .. 10, 12
Stismaticas le se wl
STRAHANI 417-8
suffusa Osha et 415-6
TENUICORNIS .. .. .. 8-9
TONNOIRI pay eealeg menage: BLY
unicolorata .. .. 10-11
WRENN cas gucah «0 sane LO
WHIPS HOIS 456 o0° +04 dy
Sarcophaga
300, 305, 334, 447, 450, 455
albiceps . .. 448, 455
alpha ROSS eee ey AS
antilope . 447-8
ANTILOPOIDES 448
aurifrons 447- 8, 452- 3
bancrofti Ai suan deta | ieee bi 15315)
beta 448-9, 453
carnaria Bee iti alte DD,
ceylonensis 452-3
communis Rael ey SAS

CHASSIMEAI eS Fat ate 2452

INDEX.

Sarcophaga crinata pig. GID)
doleschali . 448, 452
Guubxeuene Sa ene 448, 452-3
Ctaiorr ga cas, Mba errr 2
exuberans eens es V4?
fergusoni ne hice sh DD
favinenul Smee wea eo
flavipalpis .. ». 448, 452
floridensis .. .. .. 458
froggatti Bea Novel blades ee aE
fuscicauda .. 448, 454
gamma 1. 458,455
haemorr hoidalis. pais we eats:
HaLDaAxXeeo ese se a ab!
howensis as 448-9
hudsoni aie athe g pswhe Al
JAVANA crac. 2c! aw oe 448
KARNYI .. 448, 454
knabi so oa AIR, ale
knabi Re ee aw eR hate de BOC a
kohla seioeea es 452-3
lineatocollis .. .. .. 448
luzonensis Sete 452-3
marshalli hee ae eae nah 2,
milleri 455-6
misera 447- 8, 451-3
MMAtICOlOT a een e456
ochideawen ae. ow of 245)
omega ; 447, 453, 455
orientalis ee wae 4S
pallinerviste =.) ei.) ae 458
pectinata SGM ok ADO
peniculata .. Pee 4 5S
peregrina 447- 8, 451, 453-5
sarracenoides aa 452
scopariiformis eel Avera D2
securifer Roma vere LAAT,
shermani oe LOTR dein: 7
synia tm Unie Feet PAS
taenionata .. 448
tryoni 450- iL, 453
tuberosa . 447, 452
zeta Seon pts th aarti BS)

Scaevola : . 472, 484
Hookeri Meni Ser hte OD

Scamboneura 59-60
plumbea 59-60

Schizomeria Ree GAA) Le Weel? ()

Scleroderma bovistoides 257
chilense PAA i OO Tf
COLUMN uae dee Lehre
leptodermum ie cae Asi |
olivaceum Bra ae) SRD
phaeotrichum 5 Om) BGI EEADI
radicatum Sie DUNES ee neta TT
SOUOMOSUT of oo. oo Dt

Selago corymbosa .... liii

Selliera ad eer erne tas. ANE aLALT 2,

SEMES WAU AGueee nee 339
AUSTRALIS " 340- il
ING/NAKCONINGIS! 65° oo | oo axial
HY ALIPENNIS 341-2
NIGRICORNISW einen Oa
NITIDIVENTRIS nid eiyee OAL

xelii.

SEMISUTURIA PAHANGEN-
SIS ae a ts Wee 4a
TRIANGULIFERA : 341-2
SEMMNMOUTS co oo o0 oa ioe
Senecio lautus .. .. .. 195
pectinatus aR Sore tat aL
Senostoma Be he SOD
Sepimentum RA iba ett On lee)
HIRTICEPS ee he TS ONS NS
Setaria tic, CRN ote bh rand Bee 3)
BUCHANANI i 185-6
@ARINETS posh) Waseem Oe Gils.
VAG CIS een ee ete eee 1
Sherardia arvensis .. 489
SHEFVIOGSSE go 66) oo) oo. AE
WENO, oo 66 oo bo. 246
MELBOURNENSIS .. .. 445
Sigmatomera 5 oo BL
Signeta Sieh i aed hes PAPA
ntymbophonae. ee seen.
Siphonaptera XXV
Siphunculina ; 435-6
breviseta Se Pas 43 G
SLUMS eye Malek niece ee RA
Solenobaris we oO
CRYPTORHYNC HOIDES 50 a
edentata Lise erent VOLO,
Spaniopsis Dav Seating PON XOX
Spermothamnion .. 460
Sphagnum xlix, 194- 5
Spirogyra .. .. ba Che
Spongoclonium "460-2 , 469
angustatum Pais 463, 467
brachygonum aa sor 46
brounianum .. 463-6
conspicuum ae 460-3
? crispulum . 463, 469
dasyurum . 463, 466
?? debile . 463, 469
fasciculatum 461, 463-6
formosum ee iictotl | Lone ol
INTO, 55 soo oo oo CGS
latissimum . 463, 467
paradoxum . 463, 466
pastorale Nt Ue caus TER EL OL
plumigerum .. ee iO
scoparium i 461, 463, 468
scopula . 463, 468
violaceum .. 463, 468
wilsonianum 461, 463, 465-6
wollastonianum 463, 465-6
Stackhousia monogyna .. 199
pulvinaris ; 416-7
Steganopsis . 399, 400
melanogaster ae, ret OO
Stegomyia aegypti .. xv, xix
Stellaria flaccida .. .. 195
pungens 195, 197-8, 203
Stenopogon Be aE E ha Ms ithe tc)
Stereoborus EEE Ped eRe BONG
Stenmingomiyayeene ea OO:
Stibadocera Ser ee eee MG!
metallica Ste nays ROL
Stibadocerella... 9.5 =. 61

XCiv.

Stibadocerodes!.. 2. =. 61
australiensis acer aes (GEL
tasmaniensis BR ok Be FOL:

Stigmodera balteata 5 LAAT
bonvouloiri .. .. .. 225
elancula Bion ai Gata) Gath ead)
COMUNE, so bo Go. ao. AAAS
DEUQUETI 225-6
DILATATA Naame ie 226-7
dimidiata Reet NPE High OPN)
DUCALIS Swine 223-4
flavo-picta ie eS
PLAVO-VIRIDIS A EE ONT:
FOSSORIA eae 226-7
horni Bil tthe eed ete oO
kershawi Be ay LeU O
MIRANDA rege: 224-5
Manila «2 Wes eeec26
oleata SO evel, ns, meee
princeps Berets tre Nba ran)
recta et Oe eee oth
Tectipenmis; =. 2. 2.) 225
Sexplaciataye ey yeaa 2o
SPENCE ie) sn ee
ICANT ss | cask (Aten geEeeaO
GTHCOLOTiapsiek eae Dee eee 2
UIPIRONNIB, 66 ou oe Be
unimaculata so. oo PKS
Wilko oo oc 228

Stilbomyia 342, 344. 5
GOStANISH ae Lent on hea oa:
opulenta mle Rees sept A hy

Stilpnogaster . 387, 391

Stomatorhinia 5 A), Bax
australis ayey ese 333
cribellata Seen en See a4:
GISCOlOTA ee ee 4
hunaitas Sse a RE R39
VATE) Cee 333
PALLIDA " 339- 4
subapicalis 332-4
xanthogaster so oo Bet

Stomoxys calcitrans ae EXON

SRO HII So 6g aq. Jhhy

Stylidium graminifolium

194-5, 197-9
lineare : 204-6, 209

Styphelia longifolia .. 495

SIRAMIMEONMIBAB, 55 wa of BIR
terrae-reginae Pa Beye OO

Swainsonia coronillifolia 379

Maemopteris; 22. weeeeewl0s8
Tanyderus vel Met eos

pictus cel, Pei die verse ay
any PLemnld pic p ie.) eens
Tanyptera siawen,. somneese eon:

fumipennis .. .. 57
Tapeigaster BRUNNEIFR ONS 16
TVATACULOCELAN 2 aes mee

dens-leonis
194-5, 197-8, 217
Tasiocera.§.. 3. oe CARL

Taraxacum

INDEX.

Telesto caecilius 66. ie AG

Telicota Ben ead Rails nian OS)

Telopea Se | cothe enamine a: COLI T
speciosissima Biter ses) JL

Temnocephala Lecter ete MLV

Tetratheca thymifolia .. 379

Teucholabis 54-5, 63
Thamnocarpus .. . 460
Thaumastoptera Nar OO

Galiceatas.) 0 a0 Re eS

Themognatha bonvouloiri 225

DUCALIS 223-4
MIRANDA ‘ 224-5
princeps nee see. CAS)
rectipennis .. .. .. 225
Thereutria 395-6
amaricus oe eee Lee OWO
luctuosa BO eno eh Sola, GOO
DWI, oo ao co oo aXe
UKM 646 oo oo BOO
Thinnfeldia Hy or POS
Thrypticomyia 64, 67
doddi see Ste AY OT
Thylacodes decussatus xlvi
Thylacoleo carnifex xlvi
Thyridula go o6 2a, 440)
atroapicata .. .. .. 441
BRUNNEIFRONS ee 441-2
centralis 441-2
TUL OS Ai ei vets maw a ee 4:4)
‘Tiliqua scincoides xliii,
dhhy, 2H, Wes, ~ axDil,
499, 500, 520, 524-5,

537, 542, 546, oe 550, 553

Timareta ns : 55 OI)
(RInemylay .. Mee see uD 0)
Tipula 57-9
omissinervis Seis Uuentateecas)
Tipulodina oon ee BRS
Todea lai) Se G2 HO,
barbara .. ae 181-2
SraSCTIv aie) Nee ee eels
ANNINEAUIS bo. oo oa «oo adil
Tonnoiromyia tasmani-
CNISIS! ice eth eLOS:
Toxidia 275-6
crypsigramma Hey erh Bash PANEL
malindeva .. .. .. 277
DELON AIG os Bese tere onKt
Sexcouttata .500 See ea eenee
Toxorhina 63, 69
magna 68-9

Tracheomyia macropi .. xxv
Trachysaurus rugosus 499, 500

TPAD EZULESirie ace aR CHILD
eliena ee ofaeeeee2i6
eliena eliena ome a Ae
eliena monocycla 50 PAG
jacchoides an, BAe eINO
ACCHUS. 5 ..k ec eee ecu
DPoizalioides! ..) ee. eee eo

Trentepohlia A OR-O

Trichocera cee MATS AO
annulata Pie ree oes A

Trichodolichopeza .. .. 59

AVENOMOINOMIS 55 oo ao Hal
Trichoneura 4 55, 62
Vul Saris. - 0) ee eee)
Trichopsidea) —2e) ea aseeocH
Tricimba >. 435,442
carinata.. . 442, 444
CARINIFACIES .. 443-4
CONVEXA .. : 443-4
POLLINOSA Bt Po oc, 414183
scutellata 443-4
SIMILATA 8 443-4
Tricyphona 47, 49, 50
brevifurcata .. .. 49
constans 5.5) 1) Se me ()
formosana’ 5. eee
gmundensis .. .. .. 50
INCOMES o5 cs so BY
littoralis oo DERE EEG
novae-zelandiae .. .. 46
OOO, 65 560 co oo BO
opaca el ree 0
protea te CHO, 4)
rainieria oS
Schineriaa = ts) eee 0)
tipulinay a). 250 ee ee)
Trigonometopsis 400, 412
binotata : 5 aula
Trigonometopus . 400, 402
fuscifrons .. 402
‘Pramicra: \.3 > 4 e= eee 0
Triogma-: . 30. eee Ol!
EXCUlD ENA An ee ene ee OL
kuwanai Pant tics weal: (0)
trisulcata Sa Sas Wee Oe
Troides priamus i pro-
nomus é xlvii
Trypaneoides .. .. 400,418
Typha angustifolia oo BS
Udotea argentea .. .. 556
orientalis sol EO OO
Ula en a Ri EDO
Ulex Europaeus’ : OS eee bg
Unionella ae, 111
Uromyces F : xl vii
Urosthenes australis oo oNlhy
Utricularia dichotoma

var. uniflora . 199, 212

Valonia confervoides .. 556

IMOVANSSL 65660 06 oa DDD
Vaucheria geminata PAS x
Velleia ice MEARIZ
Velleya montana . 198, 212

paradoxa 197-8

Vermetus decussatus . xlvi
Veronica Derwentia ;
192, 194, 198, 200
Verreuxia .. .. : . 472
Viola betonicifolia ..

195, 198, 217, 379

Wahlenbergia gracilis ..
195, 197-8,379

Warnenias 25.5 .. ' 4. 460
Weinmannia .. .. .. 120
Xenocalliphora 50 00 Baht?

INDEX.

XENOHOMONEURA ..._... 419
TESTACEA Syst? ehscn | te 4a
Xenolimnobia 54-6, 62
camerounensis .. 55-6
Xenopsylla cheopis xxv, xxvi
Xenotipula ay ie te COD

XCV.
Xerotes longifolia .. 197-8
Xipholimnobia 54-5, 62
terebrina cae UN es eel tT)
Xylotrupes Lt, Coa ADD
VAN ENNACOMONIEY 55 60 oo BO

Zelandotipula .. .. =. 58

NOTES ON AUSTRALIAN DIPTERA. No. x.

By J. R. MALLOocH.
(Communicated by Dr. HE. W. Ferguson.)

(Twelve Text-figures. )
[Read 30th March; 1927.]

In this paper I present a revised synoptic key to the species of the genus
Drosophila Fallén, with descriptions of some new species, a synopsis of the species
of the genus Homoneura van der Wulp, descriptions and records of some species
of the genus Sapromyza Fallén, and descriptions of some other acalyptrate Diptera,
most of which have been received from Dr. H. W. Ferguson.

Family Drosophilidae.
Genus MycoprosopHILa Oldenberg.

This genus is distinguished from Drosophila by the extremely short anterior
pair of postsutural dorsocentral bristles, which, like the basal pair of scutellar
bristles, are almost indistinguishable and by the humped up thoracic dorsum.
All the known species are very similar in colour, glossy black on dorsum, with
yellow abdominal markings and pale yellow on venter and sides of thorax. The
lies occur commonly on the undersides of fungi in which the larvae feed.

I have before me a series of specimens belonging to a species which is evidently
undescribed and one injured specimen which may possibly belong to another
species.

MYCODROSOPHILA ARGENTIFRONS 0. SD.

Male and female—Head black, yellow on centre of frons from ocelli to
anterior margin and on lower half of occiput, the face brownish, bases of antennae
yellowish; orbits shining, rest of frons dull, when seen from in front densely
silvery; proboscis yellow below, fuscous above; palpi fuscous. Thorax glossy
black on dorsum, with a variable amount of yellow on anterior margin, sometimes
in the form of two rudimentary vittae; scutellum when seen from in front brownish
or yellowish dusted apically; pleura pale yellow, black on upper margin; post-
notum black. Abdomen pale yellow, with a black fascia across hind margin of
each tergite which is carried forward on the median line on tergites 3 to 5 and
connects with a similar fascia on anterior margin leaving two yellow spots on each,
the lateral margins of tergites wholly black. Legs pale yellow. Wings hyaline,
with a deep black mark on costa before apex of first vein and a faint cloud
extending from it over cell to second vein. Halteres black.

Head a little broader than thorax; frons at vertex nearly twice as wide as
length at centre; all bristles, except the anterior reclinate one on each orbit, long;
eyes almost bare; facial carina widened and flattened below; clypeus convex.
Thorax quite conspicuously convex, the dorsal surface rather densely fine haired.
Third costal division over half as long as second.

Length, 2-3 mm.

Type male, allotype, and 5 paratypes, Coramba, N.S.W., 15.2.25.
A

2 NOTES ON AUSTRALIAN DIPTERA, X.

Genus LEUCOPHENGA Mik.
LEUCOPHENGA MINUTA DN. SP.

Male.—Head testaceous, occiput and face, except sides, infuscated, frons
slightly darkened above, but all these parts with dense white dusting so that
in some positions they appear pale; palpi and antennae testaceous; cephalic
bristles, except on vibrissal angles, yellow. Thorax testaceous yellow, densely
white dusted, almost silvery when seen from certain angles, with evidences of a
rufous vitta along each side of dorsum; a brownish vitta along centre of pleura;
scutellum testaceous yellow; postnotum brown. Abdomen brownish, when seen
from in front with quite dense silvery dusting on dorsum. Thoracic and
abdominal hairs and bristles luteous. Legs testaceous yellow. Wings hyaline, a
narrow fuscous cloud extending from apex of posterior basal cell obliquely to apex
of first vein. Halteres missing in type.

Frons about twice as long as wide, parallel-sided, all bristles well developed;
face slightly carinate on upper half; antennae normal; cheek linear; vibrissae
short; palpi slender. Both sternopleurals long. Legs slender, no outstanding
setulae on mid tibiae. First posterior cell of wing not narrowed apically; third
costal section over half as long as second.

Length, 2 mm.

Type, Cairns District, Queensland (Dodd).

Distinguished from other Australian species known to me by the uniformly
silvery white dusted dorsum, markings of the wings, and its small size.

Genus DrosopHiLa Fallén.

As I have described a number of species of this genus since I published my
key in the first paper of this series (These PROCEEDINGS, 1923, p. 615) I deem it
expedient to present now an enlarged synopsis which includes 22 species, all at
present known to me as occurring in Australia. I have found it necessary to draw
up this key for my own convenience in identifying material and hope that it may
prove as useful to other students of the group as it has to me for that purpose.
There must be many Australian species of the genus unknown to me. Most of those
herein included are similar to the general run of species of the genus throughout
its range, but some, and especially nicholsoni, are aberrant, though not entitled,
in my opinion, to subgeneric segregation. I have not attempted to figure the
genitalia of either sex, some of which possess striking specific features, nor have I
attempted to determine the structure of the eggs of the species. Some of the eggs
are furnished with filaments at one end which vary in form and in number with
the species and they may sometimes be as readily distinguished as any of the
other stages, and in some closely allied species even more readily than the adults.

Key to the Species.
1. Wings with conspicuous fuscous markings in addition to those over the cross veins

OLiatlapex OL Ursticostale divAsion! Ara ame acke eee i Ore lstorede oe iene rere ae 2
Wings without fuscous markings except sometimes faint clouds over the cross
veins or at apex Of, MITSE) COSLALIGIVISION ec eteriellieckdiceie ot ek teiee nei ncnen mer tet 3

2. Wing with a large brownish or fuscous spot at apex of second vein, the dark cloud
at tip consisting of a brownish suffusion along the apices of third and fourth
veins, more or less coalescent in first posterior cell; mesonotum dark brown, with
three linear yellow vittae, the median one not reaching anterior margin;
scutellum yellow in centre, dark brown on sides of disc; pleura whitish-yellow,
contrasting sharply with the dark brown mesonotum .... mycctophaga Malloch.

10.

It,

12.

13.

14.

BY J. R. MALLOCH. 3

Wing without a dark spot at apex of second vein, the dark cloud at tip consisting of
a broad curved brown patch which extends from middle of third section of costa
to just over third vein and over disc of wing to beyond fourth vein, but leaves
a hyaline spot in apex of first posterior cell; mesonotum brown, with two poorly
defined paler vittae which are carried over lateral margins of scutellum, the
centre of latter brown; pleura not noticeably paler than mesonotum .........,.
SHccage Sorey sab anatis gohehan oiataretrinetotauen tebe wobacdnDoDoaDDODODCODDDCODOCS. MOOT MEW er

Fore femora with a comb-like series of microscopic setulae on apical third or more
offanteroventral surftacevepacvne theo oot accel sverelse no etalalevelguahateneuelauatevecstetcnelic s, « 4

Fore femora without a comb-like series of microscopic setulae as above .......... 5

Fore femora with short closely placed setulae on more than the apical half of
posteroventral surfaces, the longest one, at apex, not longer than the femoral
diameter; third section of costa not less than one-third as long as second; facial
CATIMA MTN ALT OW wv cere encusuet er eet Onste eh eh ove ias rede nchatie: suse cyai teen einstabe: wistelstie setifemur Malloch.

Fore femora with four or five widely spaced bristles on the entire length of each
posteroventral surface, the longest one, at middle, as long as or longer than
the femoral diameter; third section of costa about one-fourth as long as second;
facial carina much broadened below .................. immigrans Sturtevant.

Thoracic dorsum with a minute but conspicuous dark brown or fuscous dot at base
of each hair and bristle and sometimes with additional brown or fuscous dots
on parts causing aggregations which assume the appearance of larger irregular
spots, the ground colour of thoracic dorsum never testaceous yellow ....... 6

Thoracic dorsum with or without dark or pale vittae, not copiously marked with
piliferous spots or dots, if faintly marked with piliferous dots then the ground

colounsofethoraxsismtestaccousmyellowarrsen orc ook Eine eileen 9
Third costal division of wing almost as long as second; outer cross vein about one-
third as long as apical section of fifth vein ............ poecilothorax Malloch.
Third costal section of wing not over one-third as long as second; outer cross vein
much more than one-third as long as apical section of fifth vein ............ a
Facial carina practically absent except between bases of antennae, where it is sharp
andwlineanr- eyes valmostibarces fair sole eyes ie, cus sievelleeteusii=, 16) 6 eueiisi eels obsoleta Malloch.
Facial carina conspicuous below, where it is rather broad and usually longitudinally
suleate; eyes with dense stiff microscopic hairs ....................20200- 8
At least the first three tergites of abdomen with a yellowish spot on each side on
the part that is) incurved on) iventer ...5..5.25..0855.-2000e0- -repleta Wollaston.
None of the abdominal tergites with yellowish ROS as above .... hydei Sturtevant.
All hairs and bristles on insect luteous; thorax not vittate ...... flavohirta Malloch.
AMllgnairseandebristleseruscoussonmplackishwmeerirre crane recieicr el eicieneieieerene cian 10

Sides of face, frontal orbits, and two narrow submedian vittae on mesonotum which
extend over its entire length and on to sides of scutellum, densely white dusted,
the whole forming two continuous white lines on a black ground, and very
cConspicuousrsternopleuralsie2ieenr eect oleeioiene clones eicieieien. albostriata Malloch.

Thoracic dorsum with or without dark vittae, not white vittate .............. 11
WNaoreee Clorrsibon GlisinbeMhy AAI sooccoccupbaoonodUndLonb OUD OUdOKOL SHO HOU OL 12
MHOLaAcCiCndorsummentirelys witholtenvittalemerri etnies bara iaieieieraie aici reir ee cioione 16

Thoracic dorsum with five dark vittae, the intervening spaces pale grey dusted; wing
with a deep black spot and two fine bristles on’ costa before apex of first vein;
third costal section almost as long as second ............ nigrovittata Malloch.

Thoracic dorsum with four or six dark vittae, the intervening spaces inconspicuously
MAleerdusted ert pas Ge «Al epee a Reet UAE toys cherokee ORE SD. ceie tere rk et 13

Arista with but one long hair above at base, no long hairs below; face without a
carina on lower portion; third costal section not over one-fourth as long as
second, and shorter than penultimate section of fourth vein; sternopleurals 3
NM eeu rare ey crea ey encvapeorscaliey sti eic iattome tan en sie Wetemcnten sisia’ «itettetianer olsen vatabee «sees MIChOlSON?, Nn. Sp.

Semeh nici eu AKioieh aaices 14

Palpi fuscous; face with a vestigial carina only on: upper part, not carinate below;
pleura with two dark vittae; third costal division over one-third as long as
second and distinctly longer than penultimate section of fourth vein; sterno-
LSU SHA eereaeswencce ct caetistyetot ose uo siascivsl ea oa eR Reerc ia ol Me meitone | anie vous) GUase Meta s buscki Coquillett.

Palpi testaceous yellow; face with a very conspicuous carina which is continued
below middle, its lower portion broadened ..............00c ce cccccsccrccers 15

4 NOTES ON AUSTRALIAN DIPTERA, X.

15. Submedian thoracic vittae narrowed at posterior extremities, but continued to the
hind margin of mesonotum, no dark central mark in front of scutellum; the two
central series of hairs located on the inner margins of the submedian dark
550 Valter ee A er UR Seat yey eR cy en One ois eSU en OST tes ence te ees EAE lativittata Malloch.

Submedian vittae not reaching posterior margin of mesonotum; a dark mark on
centre of hind margin of latter; the two central series of hairs entirely within
the pale central vitta, the next two series on edges of the dark submedian

WIC! | Val soh patie seicvenp weeps Sa tensute vena aistie vay ohana ahve) ikat su alccuraiha Wonte Devtentene beret a ewe nai csi re Wat be enigma nN. Sp.

16. Facial carina not developed except weakly on upper part between bases of
Fey oli s$ ob 02 (2 yu Meee Bree Pa aE Croce carr, UREN EG hy tot Ho APR Cn OeNe Se PP CREATEN.S tote tastes er CRA atetins 1! GG. aT

Facial carina well developed and broadened on lower part of face .............. 18

Teepe BY ony feb. cao WON 5 eb ISLerO Dy Reigtols: aeoe tie MCIORIO cuclOke Gi O-G ON IS Gee ORs crcl wala Gidc Juscithorax Malloch.
Thorax testaceous yellow, sometimes with faint dark dots at bases of the bristles
andehairs: ‘On: (GOnSUMe oc ls ereie. scsncieshe lente rsner cue ae velscs hers sosuenauads inornata Malloch.

18: Thorax morejor less shining pla chy kore crckeuescueneney veka sicaielci aed ciel estonia tiscali ane Noten 19
Thoraxyshinine testaceousyellows wieicacccuimenercietey acess aiareteas ds IONS Rance maleate 21

19. Third costal section distinctly more than half as long as second and longer than
penultimate section of fourth vein (3:2); frons reddish or yellowish brown
except on orbits and triangle, which are shining black, the anterior reclinate

orbital bristle laterad of the proclinate one ................. sydneyensis, n. sp.
Third costal section about one-third as long as second and about as long as, or very
little longer than, penultimate section of fourth vein ...................... 20

20. Frons dark brown, the orbits and triangle dull black; anterior reclinate orbital
bristle well above the proclinate one; thorax shining, but not glossy ..........

MPEP Ie aria citi, LI SCO pee ce hence era NR LORU MUS ctrciie Tie neaiotenn ter stiey suis pie ee eicireweitopier eran ee ae sare subnitida n. sp.

Frons black, the orbits and triangle shining, anterior reclinate orbital bristle but
little above level of the proclinate one; thorax glossy ...... nitidithorax, n. sp.

21. Fore metatarsus of male with a comb of short black bristles on apical half, the
second. Ssesment without a "combyeeiiiy is = jens sale ee) clens oncie = ampelophila Loew.

Fore metatarsus and second segment each with a comb of black bristles from base

fYa) Dione tha WMI GooaecuccoosuogausucUdangd oan RoODOOSoOKaobOOUS serrata, nN. Sp.

[Drosophila brunneipennis Malloch described in These PROCEEDINGS, 1923, p. 617, has
inadvertently been omitted. In the previous key loc. cit. p. 616 it is placed next to
D. melanogaster Meigen = D. ampelophila Loew.-—Ed.]

DROSOPHILA NICHOLSONI, Nn. sp.

Male and female.—Testaceous yellow, shining. Frons opaque brownish, the
orbits, sides of triangle, and anterior margin testaceous, not shining; third
antennal segment largely fuscous; palpi and proboscis testaceous. Dorsum of
thorax slightly brownish and greyish dusted with six brown vittae, the submedian
pair of moderate width and complete, the intermediate pair much narrower and
connected with the submedian pair behind suture, the sublateral pair broad,
rather indistinct and broken at suture; pleura and metanotum largely brown;
scutellum brown in centre, narrowly yellow on sides. Abdomen shining, apical
tergites slightly darkened. Legs testaceous yellow. Wings greyish hyaline, veins
not clouded. MHalteres yellow.

Anterior reclinate bristle about one-third as long as posterior one and almost
in transverse line with the proclinate one; postvertical bristles long interfrontalia
with a few hairs anteriorly; facial carina distinguishable only on upper part;
arista with one long hair on upper side at base and some short pubescence beyond,
which is only visible under a high magnification; vibrissa single. Thorax with
two humerals, about eight series of intradorsocentral setulae, the submedian two
series on margins of the pale central vitta, and 3 sternopleurals. Legs normal.
Second section of costa about four times as long as third, the latter not longer than
penultimate section of fourth vein, ultimate section of fifth vein subequal to
penultimate section of fourth and about twice as long as outer cross vein.

Length, 2-5-3 mm.

BY J. R. MALLOCH. 5

Type male, allotype, and 2 paratypes, Perth, W.A., 15.11.1924 (Nicholson).

This species is readily distinguished from any other known species in the
genus either from Australia or elsewhere by the presence of but one long hair
on the upper side of the arista. This character was used as the distinguishing
feature of the American genus Cladochaeta by Coquillett, but the present species
differs from that genus in many characters and is so obviously merely an aberrant
Drosophila that I retain it in this genus, while noting that it apparently weakens
the claim of Cladochaeta to generic recognition.

DROSOPHILA NITIDITHORAX, Nn. SD.

Male.—Deep black, dorsum of thorax and of abdomen almost glossy. Face
whitish dusted and, like the cheeks, partly brownish yellow; frons except the
orbits and triangle velvety black; antennae black; palpi and proboscis dusky
testaceous yellow. Thorax without markings, the pleura not so conspicuously
shining as dorsum. Abdomen slightly yellow at base. Legs dusky testaceous
yellow, femora almost entirely fuscous. Wings greyish hyaline, veins unclouded.
Halteres brownish yellow.

Frontal bristles strong, anterior reclinate orbital half as long as posterior
one and slightly but distinctly above level of the proclinate one; interfrontalia
with quite dense short black setulose hairs; facial carina nose-like, rounded above
on the lower part; rays of arista about 4+ 2; vibrissae strong. Intradorsocentral
setulae in 8-10 series; sternopleurals 3. Legs normal. Third costal section not
longer than penultimate section of fourth vein and about one-third as long as
second section; outer cross vein at about 1:5 its own length from apex of fifth
vein.

Length, 2-5 mm.

Type and paratype, Perth, W.A., 15.11.1924 (Nicholson).

A robust black species which is most closely related to subnitida deseeipea
below.

DROSOPHILA SUBNITIDA, Nl. SD.

Female.—Distinguished from the preceding species by the brown colour of
the frons, the opaque frontal orbits and less shining dorsum of thorax.

There are no outstanding structural distinctions, but the anterior reclinate
bristle is larger and farther from the proclinate one, the third section of costa is a
little longer and the insect is less robust.

Length, 2 mm.

Type, Sydney, N.S.W., 6.1.25.

This species has somewhat the appearance of fuscithorax Malloch, but the
latter has no carina on lower part of face and differs in other respects.

DROSOPHILA SYDNEYENSIS, Nn. Sp.

Female.—Black, dorsum of thorax and abdomen almost glossy. Frons brownish
red, orbits and triangle shining black; face yellowish on sides, slightly white
dusted; cheeks yellowish; antennae yellowish or rufous, third segment fuscous;
palpi and proboscis yellowish. Thorax as in nitidithorax, but the pleura as
conspicuously shining as mesonotum. Abdomen with bases of basal two or three
tergites yellowish. Legs as in nitidithoraz. Wings hyaline. Halteres yellow

Frons with a few hairs in front; anterior reclinate orbital bristle short, in
transverse line with proclinate one; facial carina well developed and nose-like.

6 NOTES ON AUSTRALIAN DIPTERA, X.

Thorax as in subnitida. Legs normal. Wing venation differing-from that of
nitidithorax and subnitida in having the third costal division well over half as
long as second, usually about, or even full, three-fourths as long as it, and twice
as long as penultimate section of fourth vein, the latter subequal to ultimate
section of fifth, and about twice as long as outer cross vein.

Length, 2 mm.

Type and three paratypes, Sydney, N.S.W., 3 and 5.1.1925, and 2.4.1925.

A less robust species than nitidithoraz.

DROSOPHILA ENIGMA, 0. SD. ;

Male and female.—Very similar to lativittata Malloch in colour and general
structure. Differs from it in being paler, the ground colour being testaceous
yellow, the dorsum of thorax greyish dusted, and with paler vittae which are
narrower, the submedian pair separated by a wider space which covers four
instead of two series of the short hairs, and the other characters of the markings
as stated in the key. The abdominal markings consist of a dark brown fascia
on hind margin of each tergite which is narrowed or interrupted in centre,
widened each side of median line and again at the lateral curvature, the fasciae
rather indistinct on the lateral incurved portions of the tergites. As in lativittata
the outer cross vein is faintly clouded and the penultimate section of fourth vein
is about half as long as ultimate.

Length, 3 mm.

Type male and allotype, Sydney, N.S.W., 22.7.23, and 23.9.24. Paratypes, two
females, Toronto, N.S.W.

DROSOPHILA SERRATA, Nn. Sp. (Text-figure 1.)

Male.—Shining fulvous yellow, very similar to ampelophila Loew which it
closely resembles in many respects. The type specimen is rather immature, but
the thorax is not vittate and the abdomen has a faint dark uninterrupted apical
fascia on each tergite. The legs are yellow and the wings yellowish hyaline with
unclouded veins. Halteres yellow. Bristles and hairs fuscous.

Anterior reclinate orbital bristle about one-third as long as posterior one and
distinctly above the level of proclinate one; eyes quite densely stiff-haired; face
distinctly carinate. Thorax with six series of intradorsocentral hairs; prescutellar
acrostichals not differentiated; both humerals strong; only two sternopleurals well
developed. Two basal segments of fore tarsus with a comb-like series of short stiff

Text-fig. 1. Drosophila serrata, two basal
segments of fore tarsi of male from in front.

black bristles on anterior side, the comb on basal segment bipartite (Fig. 1). Third
section of costa about half as long as second and a little less than twice as long as .
penultimate section of fourth vein, the latter about one-third as long as ultimate
section and subequal to ultimate section of fifth vein; outer cross vein at about
twice its own length from apex of fifth vein.

Length, 1:75 mm.

Type, Eidsvold, Queensland, 2.4.24 (Bancroft).

BY J. R. MALLOCH. 7

There are several described species of the genus with the two basal segments
of fore tarsi armed with comb-like bristles, but none of these have the armature as
in this species so far as I am aware. I described one, biarmipes, from India, but
in it the combs are confined to the apical parts of each segment and do not extend
along the whole length of the anterior surfaces as here.

Genus GiTonipES Knab.
The species described below falls most readily into Gitonides, but it differs
from the genotype in having the frons much narrower anteriorly and the first
posterior cell of the wing quite noticeably narrowed apically.

GITONIDES CONVERGENS, 0. Sp.

Head brownish testaceous; upper half of occiput fuscous, lower half white
dusted; upper extremities of frontal orbits, ocellar spot and clypeus, fuscous; face
slightly white dusted; palpi brownish, paler at apices. Thorax brownish testaceous,
dorsum when seen from in front with a broad brownish centrai vitta which is
faint in front of the suture, where it is sometimes divided centrally and which is
divided between suture and hind margin, a broad branch curving to each side
and between the two pairs of dorsocentrals, and a narrower central one continuing
to hind margin, the disc laterad of the anterior and posterior portions of vitta
white dusted, between these pale markings and lateral margins there is a broad
brownish vitta; humeral angles testaceous; scutellum when seen from in front
with a brown central line which broadens out and covers apex, a fainter brown
mark on each basal angle and a less distinct dark mark on disc each side of the
dark central line, the latter surrounded by whitish dusting. Abdomen testaceous,
all tergites except the basal two largely or entirely black. Legs testaceous, femora
browned, tibiae dark at apices, the mid pair most conspicuously so. Wings
hyaline. Halteres yellow.

Frons at vertex about one-third of the head width, much narrowed anteriorly,
at front margin not half as wide as long in centre; proclinate orbital bristle
well above middle of frons, the anterior reclinate bristle quite small and about
midway between the others; arista bare; face slightly carinate; antennae normal,
inserted at middle of profile. Thorax with two pairs of postsutural dorsocentrals,
the anterior pair short, about eight series of intradorsocentral setulae, one humeral
and two sternopleurals; scutellum convex. Abdomen broad and short. Legs stout,
normal. Inner cross vein at two-fifths from apex of discal cell; outer cross vein
at less than its own length from apex of fifth vein; ultimate section of fourth
vein over three times as long as penultimate section; first posterior cell quite
noticeably narrowed apically.

Length, 3 mm.

Type, Hidsvold, Queensland, 1924 (Bancroft).

Family Agromyzidae.
Subfamily MILIcHIINAE.
DESMOMETOPA VARIPALPIS, 0. Sp.

Female.—Head black, whitish-grey dusted, with the usual opaque black
M-shaped frontal marking; cheeks yellowish below; palpi testaceous, with
conspicuous irregular black spotting. Thorax black, slightly shining, evenly
greyish dusted and without vittae. Abdomen greasy in type, black, and probably
less dusted than thorax. Legs black, tarsi yellowish, darker at apices. Wings
hyaline. Knobs of halteres yellow.

8 NOTES ON AUSTRALIAN DIPTERA, X.

Each orbit with the two upper bristles directed outward over eyes, the two
anterior bristles incurved; arista hair-like, almost bare; palpi large, a little longer
than head, lanceolate, broader than cheek, their apices rather pointed, bristles
short. Thorax with two pairs of postsutural dorsocentral bristles. Legs normal.
Wings as in m-atrum Malloch.

Length, 2-5 mm.

Type, Bourke, N.S.W., 6.5.26.

I know no species of this genus which kas the palpi coloured and shaped as
has this one, these organs being in all others unicolorous or yellow with dark
apices.

Family Piophilidae.
PIOPHILA CONTECTA Walker.

This species I previously listed as latipes Meigen. (These ProckEpines, 1925,
p. 316). Dr. O. Duda considers that contecta Walker is a distinct species, having
two humeral bristles instead of only one, and the second segment of the fore tarsus
about 1:5 as long as wide, not about as wide as long.

Besides the specimen already recorded by me I have seen another female
specimen from Wahroonga, Sydney, N.S.W., 26.1.26.

I have before me what appears to be an undescribed genus of this family but
have only one female specimen so defer describing it meantime.

Family Sapromyzidae.
Genus SaproMyza Fallén.

In presenting my synopsis of the species of this genus (These PROCEEDINGS,
1926, p. 33) I stated that undoubtedly there were many species still unknown to me
and as evidence of this furnish descriptions of several that have been received
since I completed the key. Under the description of each of these I have included
notes which will serve to associate them with their most closely related forms in
the key, but unfortunately there must yet be many undescribed species which can
only be distinguished from those dealt with to date by a careful comparison with
the complete descriptions or with the type-specimens.

I erect one new subgenus in the present paper for the reception of a very
striking species from Sydney, but the others I retain in Sapromyza sens. str.,
though several of them are rather aberrant from the genotype.

It will be necessary to publish a full key to the species later, but whether this
duty may fall to me or another worker time will decide.

Subgenus HENDELOMYZA, n. subg.

Characters: Face concave in profile; basal segment of antenna as long as, or
longer than, second, with some fine hairs below apically; sternopleura with but
one bristle; thorax without presutural dorsoventrals. In other respects similar to
Sapromyza.

SAPROMYZA (HENDELOMYZA) TENUICORNIS, n. sp. (Text-fig. 2.)

Female.—Head fulvous yellow, shining; ocellar spot fuscous, a fuscous line
along inner margin of each orbital stripe which curves round anterior margin of
latter and becomes wider there, almost connecting with the black spot between
each antennal base and eye; some white dusting on sides and upper margin of
frons; lower central part of face fuscous; parafacials white-dusted below the black

BY J. R. MALLOCH. 9

spot; third antennal segment brown, darker apically; arista black, yellowish at
base; proboscis and palpi black. Thorax shining fulvous yellow, with two sub-
median vittae and one near each lateral margin white-dusted; pleura entirely
white-dusted; scutellum paler yellow than mesonotum. Abdomen glossy black,
yellowish on disc basally, where it is slightly grey-dusted. Legs yellow, apices of
fore femora and tibiae infuscated. Wings honey yellow. Halteres brown.

Text-fig. 2. Head of Sapromyza (Hendelomyza)
tenuicornis, from side.

Head in profile as in Figure 2; entire frons shining; anterior orbitals rather
far from lateral margins; ocellars rather weak; arista subnude; head wider than
high; proboscis stout. Thorax with but two pairs of postsutural dorsocentrals and
one pair of prescutellar acrostichals; intradorsocentral setulae in four series in
front of suture; scutellum convex, with four bristles; prosternum almost bare;
mesopleura and sternopleura each with one bristle. Abdomen stout. Fore femur
without preapical anteroventral comb; tibiae with preapical dorsal bristle. Inner
cross vein almost below apex of first vein and at middle of discal cell.

Length, 7-5 mm.

Type, Bayview, Sydney, N.S.W., 19.12.25.

A very characteristic species which might eventually be placed in a separate
genus, though I prefer to consider it as a subgebus at present.

It must be noted that magnicornis Malloch, with which tenwicornis has some
characters in common, has two sternopleural bristles and the head quite differently
shaped. This new species does not fit into any caption of my recently published
key, the only other species having the antennae longer than the head being
magnicornis.

SAPROMYZA RIPARTA, ND. SD.

Male and female—Head dull ochreous yellow; frons brownish in centre,
ocellar spot fuscous; orbital stripes densely yellowish-grey dusted; a black or brown
spot between each antenna and eye; antennae pale brown or yellowish, third
segment and arista black; proboscis yellow; palpi black. Thorax subopaque
ochreous yellow, rather densely greyish dusted, with two conspicuous brown
submedian vittae which become wider behind, and traces of two sublateral vittae
of the same colour behind suture; mesopleura and anterior part of propleura
fuscous; scutellum brownish yellow, paler at anterior lateral angles and between
apical bristles, and with two black apical spots. Abdomen ochreous, shining, with
central part of each tergite darker and the apices paler. Legs testaceous yellow,
apices of all femora, tibiae and tarsi, and bases of all tibiae, black. Wings yellowish
hyaline. Halteres yellow.

10 NOTES ON AUSTRALIAN DIPTERA, X.

Head almost normal in form, the face slightly receding below; orbital stripes
receding from eyes anteriorly; frontal bristles all distinct; third antennal segment
about 1:5 as long as wide; arista very short pubescent; proboscis stout. Thorax
with three pairs of postsutural dorsocentrals, the anterior pair short and well
behind suture; mesopleura with one bristle; sternopleura with two bristles rather
close together; prosternum with a few very short hairs. Fore femur without
anteroventral comb; mid tibia with two long apical ventral bristles; all tibiae with
preapical dorsal bristle. Inner cross vein below apex of first vein and at middle
of discal cell.

‘Length, 7-7-5 mm.

Type male, allotype, and one paratype, Clyda R., Nov., 1925 (H. J. Carter).

This species will run to caption 11 in my key to the species of the genus, but
it is distinguished from magnifica Malloch by the unspotted wings and grey orbital
vittae and from all the others falling under caption 10 by the entirely opaque frons.

SAPROMYZA BREVICORNIS. 1. SD.

Male and female.—Head black; frons velvety, the triangle and orbital stripes
glossy; parafacials and sometimes the sides ef face yellowish, the former white
dusted; cheeks yellow below, black above; antennae fuscous, yellowish at bases,
arista black; palpi and proboscis black. Thorax shining fulvous yellow, some-
times darkened a little on disc of mesonotum. Abdomen glossy black, yellow
at base. Legs black, coxae, trochanters, bases of femora, of tibiae and of tarsi
of mid and hind legs, yellow. Wings brownish hyaline. Halteres yellow.

Head of normal form, the face centrally vertically convex; frontal bristles
normal, the anterior orbital distant from eye; antennae short, third segment not
much longer than wide; arista with very short pubescence. Thorax with three
pairs of postsutural dorsocentrals, the anterior short pair well behind suture;
scutellum convex; mesopleura with one bristle; sternopleura with two; prosternum
with a few hairs. Abdomen robust. Fore femur without an anteroventral comb;
mid tibia with an unequal pair of apical ventral bristles; preapical dorsal bristle
present on all tibiae. Wing venation as in riparia.

Length, 5-6 mm.

Type male, allotype, and two male and one female paratypes, Sydney, N.S.W.,
Sept,-Oct., 1925.

This species runs to caption 10 in my key, kaving the head black, with para-
facials yellow, but is readily distinguished from sciomyzina Schiner by having the
frons largely velvety instead of shining, and the third antennal segment not much
longer than wide, instead of twice as long as wide.

SAPROMYZA URBANA, Nl. SD.

Male.—Entirely testaceous yellow, shining, the frons not conspicuously shining,
and with undifferentiated orbits. Wings hyaline. Halteres yellow.

Anterior orbital bristle not much in front of middle of frons, shorter than the
ocellars; surface of frons with microscopic black hairs; antennae short; arista
broken in type so that it is not possible to determine the nature of the hairing if
any. Thorax with three pairs of postsutural dorsocentrals, the anterior pair rather
weak, but in this character the species is about intermediate between the two
groups defined in caption 9 of my key; hairs on thorax much weaker than in
unicolorata Malloch, the median two series on dorsum most conspicuous. Legs
slender, the fore femur with only two or three of the posteroventral bristles on
apical half well developed, the anteroventral preapical comb present but weak.

BY J. R. MALLOCH. 11

Inner cross vein below apex of first vein; penultimate section of fourth vein not
over half as long as ultimate section.

Length, 3:75 mm.

Type, Sydney, N.S.W., 8.10.25.

An inconspicuously coloured species with no outstanding external structural
characters. It differs from any Australian species except wnicolorata Malloch in
being entirely testaceous yellow, and is a less robust species than it, with weaker
hairing, and different venation. The hypopygia are distinct but I have only the
type-specimens of each and do not care to dissect these, unless compelled to for
lack of other material later.

An error occurs in caption 22, first section, of my key. The inner cross vein
is beyond, not proximad of apex of first vein in unicolorata. This feature is
correctly reported in the description of the species.

SAPROMYZA MARIAB, n. sp. (Text-figure 3.)

Male and female.—Fulvous yellow, distinctly shining. Frons dull, the orbital
stripes and triangle shining; ocellar spot black; parafacials white dusted; antennae,
palpi, and proboscis, yellow, arista fuscous. Dorsum of abdomen largely black or
fuscous. Legs yellow, apices of fore and hind femora and tibiae, fore tarsi from
before apex of basal segment, and mid and hivd tarsi from before apex of third
segment, black. Wings luteous hyaline. Halteres yellow.

Frontal bristles all well developed, anterior orbital distant from eye; third
antennal segment fully 1-5 as long as wide, rounded at apex; arista subnude; head

Text-fig. 3. Head of Sapromyza mariae from the side.

in profile as in Figure 3. Thorax with three pairs of postsutural dorsocentrals, the
anterior pair short; mesopleural hairs stronger than usual, one or two of. them
quite bristle-like; scutellum normal. Hypopygium stout, the forceps much shorter
and stouter than in flavimana Malloch. Fore femur without preapical antero-
ventral comb; mid tibia with two long apical ventral bristles; preapical dorsal
bristle weak on hind tibia. Inner cross vein almost below apex of first vein and
close to middle of discal cell.

Length, 6-6-5 mm.

Type, female, allotype, one male and one female paratype, St. Mary’s, N.S.W.,
12.8.24.

This species will run to flavimana Malloch in my key, but the latter has the
frons entirely shining, the face flat and dull, and the fore femora more largely
black, ete.

12 NOTES ON AUSTRALIAN DIPTERA, X.

SAPROMYZA OCCIPITALIS Malloch.
Hight specimens, Sydney, N.S.W., October and November, 1925.

SAPROMYZA SCIOMYZINA Schiner.
One specimen, Sydney, N.S.W., 25.10.25.

SAPROMYZA BRUNNEOVITTATA Malloch.
Two specimens, Sydney, N.S.W., 18.10.25.

SAPROMYZA STIGMATICA Malloch.
One specimen, Millgrove, V., Dec., 1925 (F. E. Wilson).

SAPROMYZA MACULITHORAX Malloch.
Four specimens, Sydney, N.S.W., July, August, and November, 1924-25.

SAPROMYZA VARIVENTRIS Malloch.

A male of this species is darker than the type female, the spot at base of
antennae on the parafacials being black, and the pale parts of the legs more
brownish yellow. The thoracic vittae are also broader, and the abdominal markings
are less regular.

Locality, Nevertire, N.S.W., 25.3.26, the same day and month as type, but a
year later.

Genus HoMONEURA van der Wulp.

In one of my recent papers (These PROCEEDINGS, 1926, p. 551) I stated that
I would present in my next paper .a synopsis of the Australian species of this
genus known to me, and though I am confidednt that my material contains but a
small portion of the native species I feel that I ought to provide a synopsis of those
already described in the interests of students of the family. This synopsis contains
only 14 species, some of them so closely allied that their separation as distinct
species has been based largely upon the structure of the male hypopygia, figures
of some of which I present herein. I have previously referred to the importance
of the male hypopygia as distinguishing characters of the species in this family
and hope that an examination of the figured examples will create an interest on the
part of some Australian student that will result in a comprehensive study of the
family on this basis. One of the species figured exhibits an asymmetry of the
inferior forceps which is unique in my experience in this family, but it appears
to be a perfectly normal specimen in other respects.

Key to the species.
1. Wings with quite conspicuous brown or fuscous clouds or spots, the cross veins and

the japex of second vein tallways Clouded iti. sievelsiohelclelin ai ie) +) -edetcel +) ished titel-(-1 2
Wings clear or yellowish, at most the cross veins slightly clouded, none of the
LONLILUIGINAL Veins ClOUGER Atta DICES micah hickaicLolonieyaasiehel Malet p Rohe Mn Meter hels 7

2. Thorax with only two of the three strong pairs of dorsocentrals behind the suture,
the other pair in front of suture; wing with conspicuous fuscous markings, the
most prominent consisting of a broad oblique fascia, the outer edge of which
is at apex of second vein, extending over fourth vein but not to margin of
wing, and enclosing a clear spot in first posterior cell; face largely fuscous,
white below; a black spot between each antenna and eye; arista plumose;
dorsum of thorax and abdomen largely fuscous; legs bicoloured, testaceous and
GATICE DOW Eee serayenciote ease iere toy svete entiea sls sacha wekeus ekeael oLebeuane heiedodersyedous atrogrisea Malloch.

Thorax with three strong pairs of postsutural dorsocentrals, or if there are but two
such ‘pairs there are no presuturals Present 6 oi. cue eee lee clele wieie a) sis elec) elelelele|islivje 3

10.

11.

12.

13.

14.

15.

BY J. R. MALLOCH. 13

Frons opaque fuscous; both cross veins of wings and apex of second vein with
fuscous clouds; arista short plumose .....................-. fumifrons Malloch,
Frons yellow or brownish yellow; in addition to the wing spots mentioned above there
are two other spots present, at or near apices of third and fourth veins ...... 4

Spots on apical part of wing at extreme apices of the veins; arista with very short
MUDESCEN CEM calit spony Canney ener eee a payee Mee tire e cl ti ls aaa eae ea apicinebula Malloch.
Spots on apical part of wing not all at extreme apices of veins; arista plumose... 5

Tips of palpi black; thoracic dorsum with a dark mark along inner side of humeral
callosities and two narrow dark vittae along inner margin of lines of dorso-
COIMETGABDIS Weve ha atatiau'e' aoute vor een ewemepe ay deoedahas <itabaasliad sreueue\a/ellecenelans omnes sine proximella Malloch.

Palpi entirely yellow; no dark mark along inner margin of humeral callosities .... 6

Cell between auxiliary and first veins of wing dark at apex; thoracic dorsum with
four faint, broad reddish vittae, the central pair extending on to disc of
YOON SONIDO. 1 fing. 6 on5/0.010 010 0.0/010-0-4)0 0 ULOlONb. 0 Grorcro1a a quaidicio mia claim aiolo ein preapicalis Malloch.

Cell between auxiliary and first veins of wing clear; thoracic dorsum not noticeably
VA Citi U Qu nvtisnsiice eOktercoute eet nites cme cme canst eed ie melne Ts Delran yon eared as Watian. ol tasts. cht a oiler aca barnardi Bergroth.

Halteres black; thorax and abdomen uniformly shining black; arista plumose; legs
fuscous, tibiae and tarsi testaceous yellow ........... signatifronus Kertesz.

Halteres yellow; thorax and abdomen fulvous or testaceous .................... 8

Thorax distinctly vittate, with 1+3 pairs of strong dorsocentrals ..................:
Bese ail csrerseiheskusaisal Mp see ne Ge oN ae aM MIs Eee stan di onanistissis meen ay ual feSertesni eleva eee wdanere eidsvoldensis Malloch,

Thoracic dorsum not vittate, with three pairs of postsutural dorsocentrals ...... 9

Abdomen with one or more of the tergites of apical part with a pair of round black
SPOYO SY MES Hears mes ican ievcunc is ecto Volto NOSJa Bec -tSiE: a bara RnnoR eeett na eh Oca rue ern ceria tera Mebane Ad rls elas 10
Abdomen without evident paired black spots ...................202005 sero astral il

Ocellar bristles short and weak; no dark dorsocentral vitta or series of spots on
abdomen; superior forceps of hypopygium stout, rounded at apices (Text-fig. 4)
AeAp eaaieudsd cutaeatiyan nest stiaho nnd seAraaieUataiay Madar oat suet ote Gulapoeay sretcjsaiiatiey oo taio sess inaieieat anes indecisa Malloch.
Ocellar bristles long and strong; a more or less distinct dorsocentral dark vitta
or series of spots on apical tergites of abdomen; superior forceps of hypopygium

slender vapicalllya (CRExt= fie Day i reteves se cuctecs erene auelel avail actors ialaneneyateranetels gordonii, a. sp.
Third antennal segment and apices of palpi black; arista pubescent .......... 12
Antennae and palpi entirely yellow; arista long haired ........ illingworthi, 0. sp.
GMA lS Ss Sete iis nil aires apie tema taene tebeiedeipetier shone: ReNIcU oh clone, wusllabisr eine milecetie alse counin fe ans Mendel eb ae 13
Va este wi seta t Sle AW ees Uh wear beta tbeg rab opel citer ama Neate farts rotearepantlben sta etteh ahgrousteMePa means ast-ehes iam ate 14
Preapical abdominal sternite glossy black only at apices of processes ..............
PU ee Ie ID Sr ea tl ra ECTS ToS RCE PEER ES OSTICY Gl HCI ERENT ETE TE ECE eR armata Malloch.
Preapical abdominal sternite entirely glossy black and heavily chitinized ..........
SS co DOD oOHeDO ODOM DOO OOOO OOO OOOO oe Ooo ONO OD HD Ola obo 0.019 o0 perthensis, n. sp.
Superior hypopygial forceps simple, but slightly curved, not recurved (Text-fig. 7)
BS AAR GIRL OTS 9 radia a Pe ckend ick Gitin cy CACHE OO23 Io tt Orc 0 Lo ee se iene er oipicd a ts DROME EO asymmetrica, n. sp.
Superior hypopygial forceps pronouncedly recurved .......-.....-eee eee reece 15
Apices of superior hypopygial forceps simple, inferior pair long and slender (Text-
TOAST shi e)D sero oenos0 6 IC SEONG: CHENG GhsOu 1d 615 /6.658; OSE OpE 6 ake DACHB RO CO ee ICMERe one eacdy perthensis, n. sp.
Apices of superior hypopygial forceps branched, inferior pair broad and serrated
om edges: @DRext-fety VQ))i Vace s'Sircnsasvete ede sastecccciehe ad: eyes sehevay ss aus celeay ee, armata Malloch.

HoMONEURA GORDONI, nh. sp. (Text-figure 5.)

Male.—Very similar to indecisa Malloch, differing in colour in having the

apices of some of the abdominal tergites blackened (possibly not invariably so) the
apical two or three tergites with a central vitta or series of elongated dark spots,
and the outer cross vein of wing slightly clouded.

A larger species than indecisa, with frontal bristles all strong, the hypopygial

forceps less robust, and the penultimate and ultimate sections of fourth wing vein
subequal in length. The antennae are missing in type specimen so that it is
impossible to determine if they are similar to those of indecisa.

Length, 5-5 mm.
Type, Gordonvale, N. Queensland, October (J. F. Illingworth).

14 NOTES ON AUSTRALIAN DIPTERA, X.

HoMONEURA ILLINGWORTHI, n. sp. (Text-figure 6.)

Male.—Similar to indecisa in colour, but without any paired black spots on
the apical abdominal tergites, each segment brownish or fuscous on hind margin
and the apical two or three with a faint dark dorsocentral elongate spot. Antennae
and palpi yellow; frons dull, orbits and ocellar region shining. Wings yellowish,
outer cross vein slightly clouded.

4 5 6 7

Figs. 4, 5 and 6. Superior hypopygial forcep of
Homoneura indecisa. H. gordowi, and H. illingworthi
respectively.

Text-fig. 7. Hypopygium of Homoneura asymmetrica.
a, from below; b, from the side.

All frontal bristles long and strong; arista plumose: third antennal segment
fully 1-5 as long as wide; face slightly convex in centre vertically. Hairs below
humeral bristles strong; centre of propleura with a few microscopic black hairs;
prosternum also haired; thorax with three strong pairs of postsutural dorso-
centrals and 8-10 series of intradorsocentral hairs; mesosternum quite regularly
setulose on most of its length. Superior hypopygial forceps long and slender,
tapered to a point (Text-fig. 6). Fore femur with an anteroventral comb; mid
tibia with three strong apical ventral bristles. Penultimate section of fourth vein
about three-fourths as long as ultimate section.

Female.—Similar to male, the abdominal markings more distinct.

I can see no fine hairs on the propleura but the specimen is not in good
condition, and the pinning prevents a view of the mesosternum. Apical genital
segment unspined.

Text-fig. 8. Homoneura perthensis, hypopygium from below.

Text-fig. 9. Homoneura perthensis, from the side.

Text-fig. 10. Homoneura armata, male hypopygium. a, from below, one
side incomplete; b, from the side, superior forceps only.

BY J. R. MALLOCH. 15

Length, 5-5-5 mm.

Type, Gordonvale, N. Queensland, October (J. F. Illingworth) ; allotype, Cairns
District, Queensland (Dodd).

Named in honour of Dr. J. F. Illingworth who collected the type specimen.

HoMONEURA PERTHENSIS, 0. Sp. (Text-figures 8 and 9.)

Male and female.—Very similar to armata Malloch in colour, but more shining,
and the penultimate ventral abdominal segment in female entirely, or almost
entirely, glossy black, instead of black at apex only.

Structurally the two species are very similar and, like asymmetrica, both
have an isolated outstanding bristle on each side of mesosternum in front of middle
instead of regular setulose hairs as in illingworthi. The female has the penultimate
ventral segment differently shaped from that of armata, there being a short blunt
process on each side at apex, while in armata the apical margin has a deep
V-shaped ventral cleft dividing the two tapered apical lateral angles; the apical
segment of the female in armata has about half a dozen stout black thorns, while
there are only fine hairs present in perthensis. The male hypopygia of the two
species are shown in Text-figures 8, 9 and 10.

Length, 5-5-5 mm.

Type male, allotype, and one female paratype, Perth, W.A., 15.11.1925
(Nicholson).

HOMONEURA ASYMMETRICA, n. Sp. (Text-figure 7.)

Male.—A smaller and less robust species than perthensis, but very similar in
most respects. Readily distinguishable by the structure of the male hypopygium

Length, 4 mm.

Type, Perth, W.A. (J. Clark).

11

Text-fig. 11. Homoneura preapicalis, male hypopygium. a, ventral
view, one side incomplete; b, side view; c, apex of inferior forceps;
d, another form, possibly a distinct species, Sydney.

Text-fig. 12. Homonewura barnardi, male hypopygium, ventral and
lateral views.

HoMONEURA PREAPICALIS Malloch. (Text-figure 11.)
HoMONEURA BARNARDI Bergroth. (Text-figure 12.)
Sapromyza barnardi Bergroth, Stett. Ent. Zeit, 1894, p. 74.
I present figures of the male hypopygia of the above-named species to
facilitate their identification (Text-figs. 11 and 12).

16 NOTES ON AUSTRALIAN DIPTERA, X.

Family Neottiophilidae.
TAPEIGASTER BRUNNEIFRONS, Nl. Sp.

Female.—Head black; frons a rich red-brown, with a spot of white dust on each
vertical angle and another at middle against each eye; parafacials, cheeks and
face, whitish dusted, the cheeks reddish brown; antennae brown, slightly dusted;
proboscis black; palpi reddish brown. Thorax shining brownish-black, with
brownish dusting, the dorsum with two submedian vittae and the lateral margins
white dusted; pleura white dusted. Abdomen shining black, yellow at base, and
centrally on apical two tergites. Legs black, a testaceous yellow ring at base and
another at middle of each tibia, tarsi testaceous yellow. Wings yellowish hyaline.
Halteres yellow.

No distinct orbitals present, the hairs on anterior margin of frons setulose;
arista subnude. Only the prescutellar dorsocentrals and acrostichals present;
scutellum short, rounded in outline, convex, haired on disc, the basal pair of
bristles much shorter than the apical pair; sternopleura with one bristle.
Abdomen short and broad, tapered at apex. Femora of all legs stout, armed with
short stout spines on apical halves of anteroventral and posteroventral surfaces;
preapical dorsal bristle indistinct on fore and hind tibiae. First posterior cell of
wing narrowed apically.

Length, 7 mm.

Type, Newcastle, N.S.W., 7.4.26.

The darkest coloured species of the genus known to me.

THE INFLUENCE OF CERTAIN COLLOIDS UPON FERMENTATION.

By R. Greic-SmiTuH, D.Se., Macleay Bacteriologist to the Society.

(Two Text-figures.)
[Read 27th April, 1927.]

Part iv. AGAR FIBRE IN THE ALCOHOLIC FERMENTATION.

In Part ii of this series it was shown (These PROCEEDINGS, 1925, p. 345) that
agar had a somewhat irregular action. It depressed and then accelerated the
alcoholic fermentation of dextrose when present to the extent of 0:2%. With 0-:1%,
it slightly accelerated and with 0:05% the fermentation was depressed. Both in
the fluid and in the solid state it markedly accelerated the fermentation of lactose
by the high temperature organism, when present to from 0:3% to 0:6% (These
PROCEEDINGS, 1924, p. 440). It had a strongly positive action when in the fibrous
or the flocculent condition in the ammoniacal fermentation of urea (These
PROCEEDINGS, 1925, p. 350) when present to the extent of 0-4%. In the lactic
fermentation of dextrose, 0-2% had no action.

The small quantities of agar were used when it was added to the medium
prior to sterilizing the fluids. It dissolved during the steaming and separated out
in the flocculent state when the fluids cooled. Half of one per cent of agar which
was the usual amount of the other colloids taken in the experiments caused the
fluids to become very viscous and it did not seem that this condition was advisable
for the alcoholic fermentation.

Small amounts were used in the following in which the agar had been
treated with acid, washed and finally dried. In all the other experiments, the
commercial fibre had been used.

TABLE 1.—THE INFLUENCE OF AGAR UPON THE ALCOHOLIC FERMENTATIONS.
Percentage of Invert Sugar Consumed.

i
Days Ae gS aD Ss 2 3 4 5
i He 2
IN GIRVGaAT Rew pyar) ete 17:5 40-8 63-7 79-4
INNS WN 35° 56 15-1 36:9 60-4 77-4
Agar 0:1% Ah hee 15-1 36:9 62-9 81:8
Agar 0:2% LN ayatis 15-7 35:8 64-4 84-2

This bears out what had already been found with crude fibre, that small
quantities (0:05%) depressed while those a little larger at first depressed and then
accelerated (These PRocrEpINGS, 1925, p. 345, Table vi).

In all these experiments with agar and yeasts, the agar had been in the
flocculent condition; it had been added to the nutritive fluids before sterilization.
In the experiment about to be recorded, half of one per cent of agar was used and

B

18 THE INFLUENCE OF CERTAIN COLLOIDS UPON FERMENTATION.

in order to avoid a thickening of the media, the agar fibre was put into a dry
tube and steamed upon several occasions after which it was dropped into the fluid
just before seeding. The experiments with agar in the ammoniacal fermentation
of urea showed that fibrous agar was almost as efficient as floccules of agar in
accelerating the fermentation.

TABLE ii.—THE INFLUENCE OF AGAR FIBRE UPON THE ALCOHOLIC FERMENTATION,
Percentage of Invert Sugar Consumed.

|
Days Pee crs | 2 | 3 | 4 | 5
I

Control, no agar .. 16-6 38°6 60-1 77-6
Agar fibre, 0:5%

In this case the agar has shown a decided accelerating effect all through the
fermentation and makes it appear that the previous indeterminate results were
caused by the use of an insufficiency of the colloid.

Part vy. OLD oR HEATED YEAST CELLS ARE NOT STIMULATED BY FULLER’S EARTH.

It was shown in an earlier paper (These PROCEEDINGS, 1923, pp. 48, 623) that the
mineral and similar colloids were able to stimulate the cells of the high temperature
organism after they had been weakened by chill. In this paper some tests were
made to see how yeast cells that had been weakened by age would respond to the
presence of one of the most active of the mineral colloids, namely, fuller’s earth.

In the first the cells were twenty hours old, in the second the culture was
twenty days old and the third was an old culture that had been kept at laboratory
temperature for four and a half months. Films of the last showed long sausage
shaped cells woven together into clumps as well as single cells. The clumps were
broken up as much as possible by repeated passage through a capillary nozzle, a
procedure employed in all cases where a homogeneous suspension is desired. The
yeast was a stock Scotch distillery yeast and the three tests are grouped together
for convenience, although they were made at intervals of three and of twenty days.

TABLE i.—THE INFLUENCE OF FULLER’S EARTH UPON THE FERMENTATION OF
DEXTROSE BY YEAST CELLS OF DIFFERENT AGES.

Dextrose Consumed Per Cent.

Age of Cells. TDA S) ads cts L | 2 | 3 | 4 | 5
a AS: — ed ea EPA 2
ZOMNOUTS mericiee Controleeces- ie 5:9 23°7 46-0 71-3 —
Fuller’s earth .. 6-7 39-9 79-9 90-6 —
Z0 Gaystearnanes Controliimre sic | 1-9 18-4 42-1 70:7 -
Fuller’s earth .. 1-7 31:3 68-7 90-0 —
PS 5udaysur. eer Controler-:s sce = 77 28°3 51-0 71-2
Fuller’s earth — 15-2 55-2 84-8 90-5

BY R. GREIG-SMITH. 19

The graphs of the three tests show that the younger cells began to ferment
quicker but once fermentation had begun there was a similarity in the curves.

The mineral colloid did not show any differentiation between a young and an
old culture of cells.

OE <i ek@Sies Con SOME

CONTROW &— — & —- -=—-—
TeEsT

Upon finding that the colloid did not have an effect upon cells weakened by
age and by contact with their own byproducts, an attempt was made to see if the
application of heat would weaken them sufficiently to enable the colloid to
exhibit an effect.

Suspensions of the same yeast were heated for half an hour at 45° and at 50°
and sown in a dextrose medium which was incubated at 28° as in the previous
cases. The following results were obtained; those for the unheated cells have
already been recorded in Table i (20 hours).

TABLE ii.m—THE INFLUENCE OF FULLER’S HARTH UPON HEATED YEAST CELLS.

Percentage of Dextrose Consumed.

Days sid feces ete 1 2 3 : 4

|
Heated at 45°

Control a 5:9 24:7 48-2 70-1

Fuller’s earth .. 5:6 1-3 81-1 90-5
Heated at 50°

Control CA ee 8 none 10:0 32:7 60:7

Fuller’s earth .. .. none 13-2 49-7 | 82-9

20 THE INFLUENCE OF CERTAIN COLLOIDS UPON FERMENTATION.

Heating the yeast at 45° had no effect upon it, the dextrose consumed in the
presence and absence of the mineral colloid was the same as by the unheated yeast
(Table i, 20 hours). At 50°, it was weakened or partly destroyed and was slower
to start the fermentation but the colloid did not accelerate the presumably
weakened cells, indeed the differences between the control and the colloid deter-
minations are less than with the unheated cells.

Heating the yeast cells therefore does not weaken them so much as to enable
the colloid to exhibit a greater differentiation.

Conclusions.—The mineral colloid, fuller’s earth, does not accelerate the
fermentation of dextrose by yeast cells that have been weakened ‘by age or by heat.

Part vi. THE NON-ADSORPTION OF THE INVERTASE OF HEATED YEAST BY
FULLER’S EARTH.

In Part v of this series, it was shown that the heating of yeast cells to 50° did
not affect them to an extent to enable the mineral colloid, fuller’s earth to show
the stimulating effect which had been observed with weakened bacterial cells.
Apparently the dextrose-fermenting function had not been affected. While this
seems to apply to the alcoholic fermentation it might not apply to the other
functions of the cell, for example, the secretion of invertase might be weakened
and it might respond to the accelerating influence of a mineral colloid. With this
idea in view, a number of experiments were made with yeast cells that had been
heated to near the lethal temperature before being grown in solutions of
saccharose.

The nutrient solution generally contained peptone 1 g., meat-extract 1 @g.,
monopotassium phosphate 0-7 g., dipotassium phosphate 0:3 g., yeast-water 100 c.c.,
and magnesium sulphate anhyd. 0-3 g. in 500 c.c. of tap-water. The solution had a
pH value of 6-8. To the above an equal volume of 16% saccharose solution was
added generally just before seeding with the heated yeast. This solution of sugar
had been filtered through porcelain to eliminate the spores of Bac. levaniformans
which are always in sugar. The test flask or flasks received 0:5% of fuller’s earth
which had been sometimes ignited, treated with hydrochloric acid washed and
dried or sometimes simply heated at 200°.

The first experiment gave a curious result and caused one to think that a
novelty had been encountered but the succeeding tests were different and were
more or less uniform. Five tests were made with heated yeast and five with
normal unheated distillery yeast. All the experiments were made at different
times and somewhat alternately, three heated, two normal, two heated and three
normal but for convenience the results with the normal cells are tabulated first.
In some of the tests the nutrient solutions were contained in small flasks and in
these the fermentation proceeded rather briskly on account of the shallow layer
of liquid formed by 50 ¢.c. The whole contents of these small flasks were taken
for the determination of the sugars. In the majority of the tests, large flasks were
used and portions of the 500 c.c. were daily abstracted for the determination of
the reducing sugars before and after inversion at 70° with 10% hydrochloric acid.
The reducing sugars were calculated to saccharose and expressed as percentages
of the saccharose originally taken. The total inversion is the sum of the inverted
saccharose present plus the saccharose lost, it being assumed that the saccharose

BY R. GREIG-SMITH. 21

must have been first inverted before being fermented to alcohol, carbon dioxide
and acids."

In heating the yeasts the cells were suspended in a nutrient fluid generally
a modified Hansen’s fluid contained in Freudenreich flasks. These were immersed
in water at 55° for varying times. After treatment, the flasks were incubated
and that with the fewest colonies, as in @, was taken for seeding the test flasks.
In 0b, the cells were suspended in a fluid of pH 6-2 and all flasks showed a good
growth even when heated for 105 minutes; the last was distributed in a fluid of
pH 5:6 and heated at 56°. This destroyed most of the cells, the 15-minute flasks
contained two colonies which were grown, distributed and used. In ec, the cells,
suspended in a fluid of pH value 5-6, had been heated at 55° for 105 minutes.
In d, the suspension medium had a pH value of 6:4 and the cells were heated for
an hour at 54°. The growth that developed consisted of a large flocculent colony
and many small ones; the small colonies were picked out, distributed and used.
Similar colonies appeared as the result of heating a suspension in pH 5-7 for
50 minutes at 54°. The large flocculent colonies were picked out and used in e.

In the normal unheated set, 1 and 2 were made in small flasks and these
show a rapid fermentation. The yeast in 5 was the crop from a single cell of
the stock yeast which had been used in all the other tests.

The speed of the fermentation varied in the tests. This was partly due to
differences in the amount of yeast added in seeding, partly to the amount of fluid
in the fermentation flasks and possibly the amount of invert sugar originally in the
fluids as the result of steaming the solution of saccharose might have had an
influence. The invert sugar present at the start of each experiment was not always
determined but in some cases it was. In d, there was 13%, e had 2%, 1 and 2 had
less than 15%, 3 had none and 5 had 4%. Each pair of the test and control fluids
was made at the same time with the same constituents, sown with the same
number of drops of the same suspension of yeast, was contained in flasks of the
same shape and capacity and was incubated side by side. One experiment may
differ from another but each control can be compared with its test.

During fermentation, the pH values of the fluids fell, that is, the acidity rose
from 6-8 to about 4:3 when from 80 to 90% of the sugar had been consumed. The
control and the test were usually the same although in a few cases there was a
difference of 0-1 or 0-2; the more fermented fluid was the more acid.

An examination of Tables i and ii and better, perhaps, of, the graphs, shows
that with the normal yeast the total inversion is slowed by the colloid, the
consumption is slowed in four cases out of five; with the heated yeast, the total
inversion was scarcely influenced by the colloid, the consumption was hastened
in all cases.

It seems to be clear that normal yeast, if we may take this distillery yeast as
a type, has its activity upon saccharose inhibited by the presence of a mineral
colloid such as fuller’s earth. This is probably the result of the adsorption of the
yeast invertase by the colloid. Upon adsorption, the enzyme is rendered inert as
has been shown by other workers and by the author (These Procrepines, 1925,

1In Part ii (1925) of this series the invert sugar as found has been given. It would
have been better to have given the total inversion as this appears to be the proper way
to express the enzymic activity. When the results of Table ii (of Part ii) are expressed
in terms of the total inversion, it is shown that asbestos behaved like the fuller’s earth
in this paper (test a) and the loss of saccharose was much the same in the absence
or presence of asbestos.

&

22 THE INFLUENCE OF CERTAIN COLLOIDS UPON FERMENTATION.

TABLE i.—THE ACTION OF NORMAL UNHEATED YEAST UPON SACCHAROSE IN
PRESENCE OF FULLER'S HIARTH.
In Terms of Original Saccharose.

Test BMGs: biatch Sve il 2 3 4 5

Days Sai cere Lie oa datncr uted 2 il 2 1 2 3 il 2 3 iL 2 3 4
Control.

Saccharose inverted ..| 46 48 } 44 45 | 21 50 30] 33 56 34 | 20 66 65 47

Saccharose consumed ..; 16 47113 54 | 24 42 69 6 34 62 81. Bl Hz

Totalinversion.. .-| 62 95 | 57 99 | 45 92 99 | 39 90 96 | 28 81 96 99

Fuller’s Earth.
Saccharose inverted ..| 32 56 | 36 44 |15 42 37 | 21 42 27 | 17 60 70 60
Saccharose consumed ..| 13 35 |12 51 |] 24 41 59 wey. Zab ay ales 40 210)

AMV EAM o5) | fol] 4 Bal | 4k Bt ae) BB OO | BO Oe). OS | BO YS OG OO

TABLE ii.mTHE ACTION OF HEATED YEAST UPON SACCHAROSE IN
PRESENCE OF FULLER'S HARTH.
In Terms of Original Saccharose.

Test Gh Sto mack Lia's a b c d e
Days TA BSE ed 2 3 1 2 3 2 3 4 1 2 3 oft 2 3
Control.
Saccharose inverted ..| 22 62 54 | 41 59 39 145 45 19116 37 24 | 34 54 28
Saccharose consumed..| 4 20 44 ]11 32 60/30 52 80 Pe ao 7A |} Ub axe 70
AtoyeyliinyeAcsiaN 65 55] FO BY OR BY Bil OO | by Oe Oi Ab 70 8G I-42 OO O8

Fuller’s Earth.
Saccharose inverted: so ilec2 2s DIDm| on D2) eco ail ed Dien) wel) ill5 to 4 | or Oe Ac eT
Saccharose consumed .. (jal tw aha A) OR BB GB OO yy cy Of | I US SO)

Stoel thse 45 45) 23 SO Oe | by OR Oe) Bil Oe IN |) PA 78 96 1 46° BO 98
pp. 50, 342). The adsorption can be only partial as the production of reducing sugars
is lessened and not prohibited. The slower production of fermentable hexoses has
been followed by a slower alcoholic fermentation in four cases out of five, a result
which seems to be quite natural. Had there been no slowing in the formation
of the reducing sugars the colloid would have hastened the alcoholic fermentation,
judged by the consumption of the sugar, as has been shown in Parts ii, iii and v
of this series.

These experiments with heated yeast were made to see if heating the yeast
to near the lethal temperature would influence the production of invertase and the
results are in a positive direction. A change did occur in this particular yeast,
grown in the particular medium. The invertase became indifferent to the
presence of the mineral colloid, fuller’s earth, and as a consequence the alcoholic
fermentation of the inverted saccharose was accelerated by the mineral colloid.
The practical outcome was that the heated yeast behaved towards saccharose just
as an unheated yeast behaves towards dextrose or invert sugar in presence of the
colloid.

23

BY R. GREIG-SMITH.

ALSWeals (ls) JES ei]

CAST Se Se 2 s) S.

IN| ©

JESS SIA TWA VISION)

JE el WAIN SIN OO

eS a5
TOS8LNOO

24 THE INFLUENCE OF CERTAIN COLLOIDS UPON FERMENTATION.

Conclusions.—Heating the yeast to near the lethal point led to a subsequent
inversion of saccharose but the agent causing the inversion, presumably the
invertase, was indifferent to the presence of the mineral colloid. The invertase of
normal yeast is sensitive to the presence of the colloid, being partly adsorbed and
producing a lessened inversion under similar conditions. As a result of the
indifference, the alcoholic fermentation of saccharose by heated yeast is accelerated
by the mineral colloids just as the fermentation of dextrose by anormal yeast is
accelerated.

A NOTE ON REPRODUCTIVE PHENOMENA IN SOME LIZARDS.

By H. CruarrE WEEKES, B.Sc., Linnean Macleay Fellow of the
Society in Zoology.

(Three Text-figures.)
[Read 25th May, 1927.]

Contents.

J. The Placentation of Lygosoma (Hinulia) quoyi.
1. Description of Material.
2. Comparison with Chalcides tridactylus, Lygosoma (Liolepisma)
entrecasteauxi and Tiliqua scincoides.
3. Comparison with the Mammalia.
4, Theoretical Considerations.
5. Summary.
II. On the Growth of the Extra-embryonic Mesoderm in Lyyosoma (Hinulia) quoyt,
Tiliqua scincoides, Egernia whitei and Egernia striolata.
III. On the Occurrence of Corpora Lutea in Lygosoma (Hinulia) quoyi, Tiliqua
scincoides, Egeriia whitei and Egernia striolata.
IV. The Omphaloplacentation of Tiliqua scincoides.

I. THe PLACENTATION OF Lygosoma (Hinulia) quoyi.

In a preliminary note communicated to this Society, 30th March, 1927, the
author recorded the occurrence of placentation in the Scincid lizards Lygosoma
(Hinulia) quoyi, Egernia whitei and Egernia striolata and stressed the importance
of the discovery of a type of allantoplacentation in these lizards which more
closely resembles that found in the Mammalia than any hitherto recorded in a
reptile. It is proposed in the present communication to describe briefly the
allantoplacentation of L. quoyi and compare it with allantoplacentation as it
occurs among the Reptilia, Marsupialia and EHutheria and to follow at a later date
with a detailed account of the placentation of L. quoyi, E. whitei and LH. striolata.

1. DrscRIPTION OF MATERIAL.

The habitat of L. quoyi ranges from sea level to approximately 5,000 feet
above sea level. Those lizards which inhabit the highest regions are much smaller
in the adult condition than those found at sea level, and during the gestation period,
which covers approximately three months, the females of the former type carry
from three to five young, while those of the latter carry from five to nine young.
The placentae in both are identical.

Given normal climatic conditions, ovulation among the females at sea level
(Sydney, N. S. Wales) occurs during the beginning of October, whereas it is two or
three weeks later among the females at 5,000 feet above sea level. There is no
sign of albumen surrounding the egg, but a thin shell membrane is present during
the early stages of development of the embryo. However, by the time the chorio-

allantoic membrane is well established, there is comparatively little shell membrane
Cc

26 REPRODUCTIVE PHENOMENA IN LIZARDS.

left between it and the uterus. The developing embryo is dorsal in position with
regard to the parent and lies with its head directed mesially whether the embryo
be in the right or left oviduct, the embryos in the one female being approximately
at the same stage in their development. The uterus surrounds each egg as a
thick white envelope which remains as an expanded chamber on the extraction of
the egg. These chambers are connected each to each by a short narrow straplike
portion of the uterus. Before the formation of the placentae the wall of each of
these so called “incubatory chambers” is uniform in structure throughout and
consists of an outer covering of peritoneum, a coat of longitudinal and circular
muscle and a narrow mucous membrane with flattened glands embedded, bounded
by a layer of fairly large columnar epithelial cells. The chorion lies immediately
beneath the uterus and consists of an outer layer of small flat ectoderm cells and
an inner layer of tapering mesoderm cells. The allantoic stalk carrying the
allantoic vessels leaves the ventral surface of the body of the embryo a short
distance in front of the hind limbs, passes upwards around the embryo and
expands into the allantoic vesicle. The mesoderm bounding the outer surface of
the allantois fuses with that of the chorion forming the chorioallantoic membrane
in the connective tissue of which ramify the allantoic vessels. The fusion
between the allantois and the chorion gradually extends around the under surface
of the chorion until the allantois reaches the yolk-sac. As in Lygosoma
(Liolepisma) entrecasteauxi (Harrison and Weekes, 1925) the allantois does not
extend around the yolk-sac but comes into contact with the upper surface only,
the mesoderm of the allantois fusing with that of the yolk-sac.

The omphaloplacenta is insignificant and functions during the early develop-
ment of the embryo. It is composed of modified uterine epithelium and chorionic
ectoderm overlying a richly vascular yolk-sac.

The allantoplacenta is highly specialized, is established approximately two
weeks after fertilization, reaches maturity two weeks later and functions until
immediately prior to the birth of the foetus. The fixation of the embryo is brought
about by the invasion of the uterine epithelium by enlarged chorionic cells
scattered throughout the chorionic ectoderm. The development of the placental
region is not restricted, placental modifications of maternal and foetal tissues
existing for the main part over the entire area embraced by the allantois.

In the modification of the maternal tissues, the uterine epithelium undergoes
partial degeneration, the dividing cell walls disappearing and the cell nuclei
becoming pushed aside by the multiplication and expansion of the underlying
maternal capillaries, so that they come to be roughly grouped in nests between
the exposed capillaries (Text-fig. 1, B). The chorionic ectoderm consists of the
scattered and enlarged cells which serve to fix the chorioallantoic membrane in
position, and for the remainder, of a layer of flattened cells which undergo the same
partial degeneration on the expansion of the underlying allantoic capillaries as
occurs among the uterine epithelial cells. Neither the uterine epithelial cells nor
the chorionic ectoderm cells completely disappear but remain clearly definable
throughout the life of the placenta, the former functioning for food secretion and
the latter for food absorption (Text-fig. 1, B). But the function of food transition
is mainly carried on by the closely apposed maternal and foetal capillaries, which
are for the most part only separated by their endothelial walls and a thin layer
of maternal and foetal cytoplasm. This type of allantoplacentation is extremely
interesting and is so far peculiar to this species of lizard, having no parallel
among those ‘reptiles for which placentation has been recorded.

BY H. CLAIRE WEEKES. 2

29. COMPARISON WITH Chalcides tridactylus, Lygosoma entrecasteaugzi AND
Tiliqua scincoides.

Definite placentation amongst lizards has been described in Chalcides
tridactylus, Chalcides ocellatus by Giocomini (1891 and 1906 respectively); in
Tiliqua scincoides by Flynn (19238); in Lygosoma entrecasteauxi by Harrison and
Weekes (1925).

In both the classic C. tridactylus and L. entrecasteauxi the allantoplacental
area is restricted and elliptical in shape, and the uterine wall is folded into
villous ridges lined by enlarged cubical ciliated epithelial cells and containing a

oe ein

Ch. Ect.

A\L. Cap. AUL. End.

Ent. Ch. Ect.C.
B. nea eleeelC a Poe nin EHO) ADU. GAR

Text-figure 1.—The two dissimilar types of reptilian allantoplacentation. (A) section
through placental region of L. entrecasteauxi showing modification and apposition of
maternal and foetal epithelium; (B) section through placental region of L. quoyi
showing partial degeneration of maternal and foetal epithelium. All. Cap., Allantoic
Capillary; All. End., Allantoic Endoderm; Ch. Ect., Chorionic EHctoderm; Hnl. Ch. Het. C.,
Enlarged Chorionic Ectoderm Cell; Ut. Cap., Uterine Capillary; Ut. Epi., Uterine
Hpithelium; Ut. Muc., Uterine Mucosa. 4

28 REPRODUCTIVE PHENOMENA IN LIZARDS.

rich network of maternal capillaries. The chorionic ectoderm cells are columnar,
extremely elongated and in L. entrecasteauzi, ciliated. There is no interpenetration
of chorionic ectoderm into the maternal crypts, the fixation of the embryo being so
slight that the least handling in the preparation of material separates the maternal
and foetal membranes (Text-fig. 1, A). As Flynn (1923, p. 76) possessed but one
stage in the development of the allantoplacenta of J. scincoides and further, as his
account is extremely brief and without the necessary illustrations, it is impossible to
deduce correctly the type of allantoplacentation present in that lizard. Hesays (1923,
p. 76) that “the union between chorion and uterine epithelium is very intimate.
The uterine epithelium apparently consists of a single layer of very flattened cells,
while the chorionic ectoderm has proliferated greatly, is much vacuolated,
resembling a typical plasmodium, and is formed in the main of markedly enlarged
cells with large nuclei and connected together by amoeboid processes. These
processes insinuate themselves into and between the maternal cells. . . .’ Hence
it can be seen that in each of the three lizards C. tridactylus, L. entrecasteauxi and
T. scincoides the foetus obtains its food by the glandular activity of the uterine
epithelium and the absorbing and phagocytic powers of the chorionic ectoderm.
In other words it is the epithelial tissues which play the important part in food
transition (Text-fig. 1, A).

This is a distinct type of allantoplacentation and it differs from that existing
in L. quoyi since in the one, the epithelial tissues are modified to perform the
function of food transition, while in the other the epithelial tissues partially
degenerate to allow the maternal capillaries to pass maternal materials more or less
directly to the foetal capillaries (Text-fig. 1, B). The relationship which exists
between these two types of placentation is somewhat similar to that which exists
between the indeciduate and deciduate types of mammalian placentation, although
the difference which occurs between the two reptilian types is not as great as
that which occurs between the two mammalian types. The placentation of
C. tridactylus, L. entrecasteauxi and T. scincoides resembles the indeciduate form
of mammalian placentation in essentials, allowance being made for the compara-
tively simple structure of the reptilian uterus and the difference in the sizes of
the developing embryos. The placentation of L. quoyi resembles that of the three
above mentioned lizards in that there is no invasion of maternal tissue but in
other respects it resembles the deciduate mammalian type.

3. COMPARISON WITH THE MAMMALIA.

Perameles is the only marsupial for which allantoplacentation has been
recorded. The placentation of Perameles was described by Hill (1897), redescribed
by Flynn (1923), and these authors hold different opinions as to the nature of the
placenta. Hill claims (1897, p. 387) that the uterine mucosa undergoes hyper-
trophy; that the vessels in the mucosa increase in size and number; that the
uterine epithelium changes into a vascular syncytium, the nuclei becoming grouped
together in nests situated in lobular projections of the deeper surface of the
syncytium; and that the maternal corpuscles pass up between the syncytial
lobules and form a network beneath the epithelial protoplasm. He claims further
that the embryo becomes attached to the prepared maternal wall by means of
enlarged chorionic ectoderm cells, which eventually degenerate over the placental
area proper; that the allantoic capillaries now directly reach the vascular surface
of the maternal placental syncytium to which they become intimately attached,
dipping down into the depressions on its surface and forming in places a regular

BY H. CLAIRE WEEKES. 29

interlocking system, and that finally the foetal and maternal blood streams are now
only separated by their thin endothelial walls and perhaps a layer of syncytial
protoplasm (Text-fig. 2).

Flynn claims (1923, p. 175) that the chorionic ectoderm does not completely
degenerate but actively invades the maternal tissues and thus brings the type of
placentation found in Perameles in line with that found among the eutherian
mammals, more especially the Carnivora ‘where there are the same characteristics
of passivity of uterine epithelium and activity of the trophoblast with a division of
the latter into a cytoblastic and plasmodial layer’’.

PSG
MC Aen

Text-figure 2.—Section through allantoplacental region of Perameles (after Hill).
All. Cap., Allantoic Capillary; All. End., Allantoic Endoderm; Syn. Cap., Syncytial
Capillary; Syn. L., Syncytial Lobule. :

However both authors agree that the foetal membranes of Perameles are
closely apposed to the maternal wall and that a degeneration of epithelial tissues
leads to a close proximity of maternal and foetal blood streams, a condition of
affairs hitherto only met with among the Eutheria. But it will be recalled that
in L. quoyi there is the same partial degeneration of epithelial tissues and mutual
apposition of maternal and foetal blood streams, and further that this is the only
recorded case among reptiles of this type of allantoplacentation. Hence the
deduction that the allantoplacentation of L. quoyi more closely resembles that of
the Mammalia, than any hitherto recorded in a reptile.

4. THEORETICAL CONSIDERATIONS.

As this is a preliminary communication, it is unnecessary to enter upon a
lengthy discussion of the significance of the allantoplacentation of L. quoyi, yet
the type of placentation is so interesting and its bearing upon phylogenetic
questions of such importance that a brief discussion of the outstanding features
of interest is warranted.

The importance of the nature of the allantoplacentation in L. quoyi is
indicated when it is demonstrated that in the genus Lygosoma two members,
namely L. entrecasteauxri and L. quoyi, have each developed distinct types of
placentation between which there is almost as much difference as exists between the

30 REPRODUCTIVE PHENOMENA IN LIZARDS.

indeciduate and deciduate mammalian types (Text-fig. 1 A,and B). The occurrence
of these distinct types of placentation in two members of the same genus is, to my
mind, conclusive evidence of the independent evolution of the placenta in each.
Hence I now claim that besides the three lizards C. tridactylus, T. scincoides and
L. entrecasteauxi, a fourth lizard, L. quoyi, has independently developed an allanto-
placenta, a fact which further strengthens the suggestion in a previous paper on
lizard placentation by Harrison and Weekes (1925, p. 484), that “‘the occurrence of
true placentation in two not very closely related Scincid lizards Lygosoma and
Chalcides, in Australia and Europe respectively, indicates that the allantoplacenta
is a functional adaptation, which may have arisen independently many times in
evolution and upon the mere occurrence of which phylogenetic statements cannot
justifiably be based’’.

In his account of the placentation of Perameles, Flynn says (1923, p. 127),
that “according to Hill’s account the placentation of Perameles is without parallel
in the whole mammalian group” and he adds that this fact has led even Hubrecht
to suggest (1909) that the placenta of Perameles on further investigation may after
all prove to be more comparable with some one or other of the placentae of the
Eutheria. Flynn studied the placentation of Perameles with the object of
discovering, if possible, some link between it and the eutherian type and he claims
that his investigations revealed satisfactory results, the placentation of Perameles
being comparable to some extent with that found among the Carnivora. But since
two essentially dissimilar types of allantoplacentation occur in two members of
the reptilian genus Lygosoma, and further, since there is a much closer relation-
ship between these two members than exists between the marsupial and eutherian
mammals, I suggest that the fact that the placentation of Perameles as described
by Hill may have “no parallel among the whole mammalian group” need not be
as incredible as Hubrecht and Flynn suppose.

It will be recalled that in L. quoyi the chorionic ectoderm of the placental
region is inactive and partially degenerates after the fixation of the embryonic
membranes, allowing for a close proximity of maternal and foetal capillaries. Since
such a specialized type of placentation has presumably been independently evolved
in this viviparous lizard from oviparous stock as an adaptation to environment, I
venture to suggest that the somewhat similar placental condition in Perameles
(after Hill) may be the result of a similar response to environment and may be
independent of any genetic relationship with that of the Hutheria.

5. SUMMARY.

(a) Omphaloplacentation and allantoplacentation are recorded for the vivi-
parous Scincid lizard Lygosoma (Hinulia) quoyi, the type of allantoplacentation
being compared and contrasted with that of the Reptilia and the Mammalia.

(6) The occurrence of two distinct and dissimilar types of allantoplacentation
in two species of one reptilian genus indicates that the allantoplacenta is a
functional adaptation evolved independently in each species.

(c) The difference between these two types of allantoplacentation in two
closely related lizards emphasizes the fact that a similarity between the marsupial
and eutherian types of allantoplacentation need not necessarily be expected.

(d) The similarity between the independently developed allantoplacentation of
the specialized reptile L. quoyi and that of the specialized marsupial Perameles
(after Hill) suggests that a similar response to environmental conditions has
occurred in these widely separated forms.

BY H. CLAIRE WEEKES. 31

II. ON THE PECULIAR GROWTH OF THE EXTRA-EMBRYONIC MESODERM IN
Lygosoma (Hinulia) quoyi, Tiliqua scincoides, Egernia whitei
AND Hgernia striolata.

In each of the above mentioned lizards the growth of the extra-embryonic
mesoderm is normal until the formation of a temporary sinus terminalis. The
mesoderm then dips into the yolk-sac and continues its growth, not over the surface
of the latter as is normally the case, but embedded in its substance. As the
mesoderm grows into the yolk-sac it splits into two layers as it does in the normal
condition when forming the extra-embryonic coelome. Thus an outer layer of
endoderm is separated from the bulk of the yolk-sac, a condition not hitherto
_ recorded (Text-fig. 3, Y.S. End.).

Ut.Muce. UGEDL Ut-Cap. %

ChEct. Y.SEnd. Som.tMes. Spl.Mes. _Y.S.Cap. | S

Text-figure 3.—Section through omphaloplacental region of T. scincoides showing
apposition of chorionic ectoderm to maternal villous ridges. Ch. Ect., Chorionic Ectoderm ;
Som. Mes., Somatic Mesoderm; Spl. Mes., Splanchnic Mesoderm; Ut. Cap., Uterine
Capillary; Ut. Epi., Uterine Epithelium; Ut. Muc., Uterine Mucosa; Y.S., Yolk Sac; Y.S.
Cap., Yolk Sac Capillary; Y.S. End., Yolk Sac Endoderm.

III. ON THE OCCURRENCE OF CorPpoRA LUTEA IN Lygosoma (Hinulia) quoyi,
Tiliqua scincoides, Egernia whitei aANv Egernia striolata.

After ovulation the ruptured follicles in the ovaries of each of the four above
mentioned lizards become yellow in appearance and richly vascular. This is due
to the presence of a corpus luteum in each, the growth of which is rapid, each
follicle being completely filled with luteal cells in less than a week after ovulation.

IV. THE OMPHALOPLACENTA OF Tiliqua scincoides.

Flynn (1923) described the occurrence of an allantoplacenta in 7’. scincoides
but made no mention of an omphaloplacenta. The presence of an omphaloplacenta
in this lizard is now recorded (Text-fig. 3).

Both immature and mature stages in the development of the omphaloplacenta
are available, the placenta apparently functioning throughout the development of

32 REPRODUCTIVE PHENOMENA IN LIZARDS.

the embryo. The area of placental modification extends over the entire under
surface of the yolk-sac where the bounding layer of chorionic ectoderm comes
into contact with the surrounding uterine wall.

The uterine wall and the chorionic ectoderm are thrown into a series of folds
which interlock, recalling the attempt at the formation of an omphaloplacenta in
C. tridactylus and C. ocellatus (Giacomini, 1891 and 1906 respectively), and the
type of omphaloplacentation found in the marsupial Didelphys (Selenka, 1886;
Osborne, 1888). The folded wall of the uterus is thickened over the placental
region and is lined by enlarged epithelial cells containing large, deeply staining
nuclei. The chorionic ectoderm consists of extremely narrow elongated cells which
fit into the maternal crypts, there being no actual interpenetration of maternal
and foetal tissues. The arrangement of the foetal tissues surrounding the yolk-sac
is not usual owing to the peculiar growth of the extra-embryonic mesoderm
mentioned above. Beneath the chorionic ectoderm there is a thin layer of yolk-sac
endoderm containing yolk granules. Between this and the main yolk-sac is the
extra-embryonic mesoderm which has differentiated in the normal manner into
somatic and splanchnic layers. The somatic mesoderm together with the above
mentioned layer of yolk-sac endoderm and chorionic ectoderm is separated from
the yolk-sac, and it is this band of tissue which is folded to fit into the maternal
erypts (Text-fig. 3). The splanchnic mesoderm lines the yolk-sac and gives rise
to a complex network of blood vessels which ramify through the substance of the
latter.

Detailed accounts of these phenomena are in course of preparation.

References.

FLYNN, 1922.—The Phylogenetic Significance of the Marsupial Allantoplacenta. Proc.
LINN. Soc. N.S.W., xlvii, pp. 541-544.
, 1923.—The Yolk-sac and Allantoplacenta in Perameles. Quart. Journ. Micr.
Sei, ixvaiiy dl, ‘pp: 124-182"
, 1923.—On the Occurrence of a true Allantoplacenta of the Conjoint Type.in an
Australian Lizard. Rec. Aust. Mus., xiv, pp. 72-77.
GIACOMINI, 1891.—Matériaux pour l’étude du développement du Seps chalcides. Arch.
Ital. de Biol., xvi, pp. 332-359. (Reprint from Monitore Zool. Ital. Ann., ii, Nos. 9-10.)
, 1891a.—tber die Entwickelung von Seps chalcides. Anat. Anz., vi, pp. 548-551.
, 1906.—Mem. Accad. Bologna (Sez. Sci. Nat.), Ser. iii.
HARRISON and WEEKES, 1925.—On the Occurrence of Placentation in the Scincid Lizard
Lygosoma entrecasteauxi. Proc. LINN. Soc. N.S.W., 1, pp. 472-486.
HILL, 1897.—The Placentation of Perameles. Quart. Journ. Micr. Sci., xl, pp. 385-442.
OSBORNE, 1888.—The Foetal Membranes of the Marsupials; the Yolk-sac Placenta in
Didelphys. Journ. Morph., vol. 1.
SELENKA, 1886.—Studien iiber Entwickelungsgeschichte der Thiere; Das Opossum
(Didelphys virginiana).

NOTES ON AUSTRALIAN MOSQUITOES (DIPTERA, CULICIDAE).

Part i. THe ANOPHELINI OF THE MAINLAND.
By I. M. Mackerras, M.B., Ch.M., B.Sc., Linnean Macleay Fellow
of the Society in Zoology.

(Three Text-figures. )
[Read 27th April, 1927.]

The valuable revision of the Australasian Culicidae by Edwards (1924) has
placed our knowledge of the family on a sound basis and enabled Australian
workers to undertake the study of local problems with some degree of confidence.
In this series of papers it is hoped to clear up some of the taxonomic problems
which he left obscure, largely because of the inaccessibility of the Macleay types,
to discuss the status and relationships of a number of species, and to add such
fresh information as is available with regard to distribution, habits and life
histories.

Edwards’s work was soon followed up by Hill (1925), who made a very full
and detailed study of the status and distribution of the Anopheles of the
Myzomyia group with special reference to those occurring in Northern Australia
and the adjacent islands. In the present paper his work is extended, special
attention being paid to the males and larvae. The status of the various forms of
A. annulipes Walk. is discussed and the opinion expressed by Hill (1925) and
Ferguson (1926) as to the synonymy of A. mastersi Sk. with this form is
confirmed. Keys are given to the females and larvae of all species occurring
on the mainland and figures of, and notes on, the male hypopygia. The life
histories of A. atratipes Sk. and A. stigmaticus Sk. and the larva of A. amictus
Edw. are described for the first time.

A very great deal of material has been available for study, and I wish
particularly to express my indebtedness to Dr. E: W. Ferguson for advice and for
the use of material and to Mr. H. Hacker of the Queensland Museum for the
opportunity of examining the fine series of northern forms of A. annulipes Walk.,
of A. amictus Edw. and of A. corethroides Theo. in their collection, without which
this paper could not have been nearly as complete as it is.

Key to Adult Females.

1. Conspicuously ornamented species with numerous white markings on wings and

TES i CST OSS oh be EO EXeV OMT en IE -Bia Uta clo Glo’ dlo ChORGES 6-O,p SIOEOINIOND Biola ooidiora.y Slold aude cone 2

More concolorous brown or black species. (Subgenus Anopheles) ............ 6

2. Pale apical half of the antepenultimate segment of the palpi divided by a dark
Seah gVesis ami tAaT Hic. Sachi et ROT Stee aen EME enor Gio 10'S CoO Gre OS © OED OT Bt Ore ore © punctulatus Don.

Pale apical half of the palpi continuous and without such ring ................. 3

38. Proboscis pale on at least portion of the apical half ......... annulipes Walk. (part)
IP TODOSCIS/AENUIRE] VASA AT KEM is cle a lahcls, Sot ROR Oe sae SUE OU w wtial cuca esac, ail oh hap et aberensearene che 4

4, Abdominal tergites 2-7 devoid of scales ................ annulipes Walk. (part)

Abdominal tergites 2-7 with numerous flat pale scales

34 NOTES ON AUSTRALIAN MOSQUITOES, i.

5. Tarsi apically banded only .....-.-. esses eee reer eee e ee ee eter recess amictus Edw.
Tarsi apically and basally banded ...........---++-+.. amictus Edw. Type C, Hill.

6. Small brown species with the basal four-fifths of the hind femora creamy to yellow
GYNESOUONSKY gouooousocdcdodoedgnGuUDdedHoDDaD ODM GDOA DOOD OOONK. stigmaticus Sk.
Larger black species with the femora not so marked .......................-. 7

7. Palpi with bushy outstanding scales; wing with a white spot beyond the middle
of the costa; hind tarsi with narrow white rings ............ bancrofti Giles.

Palpi normal; costa without such spot; hind tarsi entirely dark .... atratipes Sk.

Key to Larvae.

1. Inner anterior clypeal hairs widely separated and closer to the outer anterior

clypeal hairs than to each other (Subgenus Myzomyia) ..........-.+.+.00-, 2

Inner anterior clypeal hairs very close together (Subgenus Anopheles) ........ 4

2. Posterior clypeal hairs closer together than the inner anterior and projecting well
LoyewronaKsl Wa) INGEKY GoadooddoocodooscoodocvaDGed dn soUODaDCo OOD GORE amictus Kdw.
Posterior clypeal hairs wider apart than the inner anterior and not reaching to the
AMIGO TANTS Oe UA MEAGC! o's oonccdconcodgandounOb GoD DCO OODOoDOUDObODO OOH 3

3. Posterior clypeal and inner occipital hairs multiple .............. annulipes Walk.
Posterior clypeal and inner occipital hairs single ................ punctulatus Don.

4. Outer anterior clypeal hair strongly branched .................... bancrofti Giles.
Omice Aiusowore Ghyreyl inevie SANS sodbooooougoobbopndobdboodoDb UGC ObouGHOBODGS 5

5. Larger black larvae with the antennal plume large and strongly branched ........
= ER erp pat LEU rales Le aimee Orn hE HORST EE eC chiiiis, BREET A CheRERE OFLO OFO-0 sd. DIA atratipes Sk.
Small brown. larvae with the antennal plume very small and finely branched ......
Sh A Oo INR tag eh at Sh Seen coh ct Mapa Yipes tat ns Sey se RMS EUG ITO TACT SMES
Note.—A. punctulatus Don. is included in the keys since it has been recorded from
the extreme north of the mainland. ‘The characters available from the descriptions for
the separation of its larva are not all that might be desired; a critical reexamination of
this species will probably reveal others of greater value. It is to be noted that the
above key to adults will not apply to males of the Myzomyia group; comparison with
known females and a critical examination of the palpal and leg markings, of the
abdominal scaling, and especially of the hypopygium are the only means of placing
specimens of this sex.

ANOPHELES (ANOPHELES) ANNULIPES Walker.

This is the common species on the mainland. Its range is from Cairns to
Tasmania and it is found in the dry inland districts and on the mountains to a
height of 5,000 feet as well as being common in the coastal area. Records from
the islands to the north of Australia require confirmation; most, if not all,
undoubtedly refer to other species of the group. In the southern part of its
range it is the only member of the subgenus met with, but in South Queensland
A. amictus Edw. appears in company with it and gradually replaces it further
north. A. annulipes Walk. bites chiefly in the evening and at night but
occasionally also in the middle of the day; its resting attitude is about 45° with
the surface. This species is a presumed malaria carrier in Australia. The
evidence for its relation to the sporadic cases which have occurred in New South
Wales is very strong, particularly so in a recent infection acquired at Narrabeen,
near Sydney (Money, 1926). In Queensland, however, the question is more
obscure, since A. annulipes Walk. and A. amictus Edw. have been confused in the
past and either or both may be carriers, while A. bancrofti Giles must also be
considered as a probable vector.

In the female sex three main types may be recognized:

1. Specimens with the proboscis entirely dark and the abdomen devoid of scales on
tergites 2-7. This is the typical annulipes Walk., and is found in Tasmania, in
the coastal districts of the mainland south of Sydney more particularly in the

cooler months, and on the mountains. It is relatively rare.

BY I. M. MACKERRAS. 35

2. Specimens with the proboscis pale on the apical half or less and with few or no
scales on tergites 2-7 of the abdomen. This is the mastersi Sk. form and is by
far the most common type throughout the entire range of the species except
possibly in the extreme south and north.

3. Specimens with the proboscis pale on the apical half and with more or less
numerous pale scales on tergites 2-7 of the abdomen. These scales are usually
narrow, but may be flat and fairly broad though always distinctly narrower
than those seen in A. amictus Edw. This form is common in the north, but
also occurs in New South Wales, where, however, it is distinctly rare.

It is to be noted that all stages linking 1 to 2 and 2 to 3 are present in the

very extensive series I have examined, but none linking 1 to 3. It appears that
the mastersi type is the continuous dominant form and that there is a tendency
to differentiate out, at the one end of the range into a type with an entirely dark
proboscis, and at the other into a type with scales on the abdominal tergites. In
the opinion of the writer this process has certainly not gone sufficiently far to
warrant giving varietal or subspecific names to the different forms.

Females of A. amictus Edw. are easily separated from any of the above three
types by possessing both an entirely dark proboscis and numerous pale flat scales
on tergites 2-7 of the abdomen.

In the case of the males the proboscis is always entirely dark or almost so;
consequently it is only possible to recognize two types, one with scales on the
abdominal tergites and one without. There is here also every stage linking
the two groups and the hypopygium is constant throughout the entire series.
Males with numerous scales on the abdominal tergites are difficult to separate
from A. amictus Edw., but may be distinguished by the fact that in A. annulipes
Walk. the scales are markedly narrower and usually less numerous and of course

by the hypopygial characters.

Text-figure 1.—a Hypopygium of A. annulipes Walk.; b Claspeties of three specimens
of same; c Claspettes of two specimens of A. amictus Edw.; d Hypopygium of
A. stigmaticus Sk.; e Hypopygium of A. atratipes Sk.; f Side-piece and its appendages of
A. bancrofti Giles.

a Cc

The side-piece of the male hypopygium (Text-fig. 1 a) bears four basai
spines usually arranged as in the figure though there is some variation. The
leaflets are four or five in number and are rather broad; they are about half the
length of the aedeagus. The claspette is rounded and there is some variation in the
arrangement of its setae. Edwards (1924) describes this structure as bearing a
long apical hair and a single accessory one which is at least half as long as the
apical one; this arrangement is frequently met with, but many specimens have
two accessory hairs half as long as the apical one, and sometimes also a third
which is a little shorter than the others. Variations are met with in the two

36 NOTES ON AUSTRALIAN MOSQUITOES, i.

sides of the one specimen. The length of at least one or two of the accessory hairs
is constantly as much as half the length of the main hair in all specimens
examined, an important point of distinction from A. amictus Edw. in which these
aecessory hairs are never more than a quarter the length of the main hair
(compare Text-fig. 1 b and c).

The larvae usually live in clear water, preferably in soakage pools and small
swamps in which there is some algal growth, but they may also be found (more
often in Queensland than in New South Wales) in rock pools, in muddy creeks
and waterholes, and even in somewhat polluted water. I have twice taken them
in running water, in the Burnett River at Hidsvold, S. Queensland, and in the
upper Hunter River, N.S.W. In both instances they were clinging, presumably by
means of their caudal grappling hooks, to filamentous algae in shallow water
where the flow was not very rapid. At Hidsvold the alga was a species of
Spirogyra and the association was almost as close as that seen in the case of
C. basicinctus Edw. and C. bitaeniorhynchus Giles, the lavae remaining submerged
for long periods attached to the alga, from which apparently they drew their
oxygen as well as their food supply. This adaptation is distinctly curious in view
of the more usual surface-living habits of the species.

(Oy za) a e
Text-figure 2.—Heads of larvae of a A. annulipes Walk.; b A. amictus Edw.; c
A. bancrofti Giles (after Cooling); d A. atratipes Sk.; e A. stigmaticus Sk.

Cooling’s (1924a, 19246) descriptions of the larva and pupa are rather
incomplete and these notes and figures are intended to amplify his. The head is
brown and has the pigment arranged in a definite and rather characteristic
pattern, not uniformly distributed as in the species of the subgenus Anopheles
described below. The antenna bears a short sublateral spine about the middle
of its length. The cephalic chaetotaxy appears to afford the only reliable characters
for the separation of the related species and is shown in Text-fig. 2 a. The inner
anterior clypeal hairs are set well apart and much nearer to the outer anterior
clypeal hairs than to each other; they sometimes bear two or three very short,
fine plumes which are hardly longer than the diameter of the hair. The outer
anterior clypeal hairs are about two-thirds the length of the inner and are
strongly plumose; in one specimen, probably as an aberration, these hairs were
strongly branched. The posterior clypeal hairs are set further apart than the
inner anterior, have four branches, are short and do not project as far as the

BY I. M. MACKERRAS. 37

anterior border of the head. The frontal and subantennal hairs are normal. The
inner occipital hair is trifid and the outer one bears a few plumes. The palmate
hairs and other characters agree with Cooling’s descriptions and figures; they do
not appear to offer any assistance in discriminating between the related species.
The pecten (in the sense used by Christophers) bears three long teeth as well as
the end ones, but is somewhat variable.

The pupal skin is brown and bears no specially pigmented areas, except for
the trumpet which is lightly powdered with pigment. The abdominal chaetotaxy
is shown in Text-fig. 3 a@ and does not require further description; its characters
are most valuable for the recognition of the pupae; the stout lateral spines
decrease in size proximally and are very short on the basal segments. The
trumpet (Text-fig. 3 d) is oblong in shape and is slightly shorter and broader
than that of A. stigmaticus Sk.

ANOPHELES (MyzoMyIA) AMICTUS Hdwards.

A larger, more yellowish species than A. annulipes Walk., from which it is to
be distinguished in the female by the combination of an entirely dark proboscis
with numerous scales on abdominal tergites 2-7, and in the male by the hypopygium.
The claspette bears two or three fine short hairs which are always less than one-
quarter the length of the main hair (Text-fig. 1 c and 0). The abdominal scales
of both sexes are broad, flat, and yellowish in colour. These characters might
be considered as very doubtfully of specific value, even though constant, and the
statement by Hill (1925) that he obtained typical females of A. annulipes Walk.
among the progeny of A. amictus Edw. females would undoubtedly lead one to
sink the latter name as a synonym, were not the question further complicated
by the discovery that the larvae are.very distinct. In view of this one cannot
but let Edwards’s name stand until the whole question can be reinvestigated and
Hill’s breeding experiments repeated and extended with special reference to the
‘males and early stages.

I have only seen females of Hill’s type “C.” This form is easily recognizable |

on this sex and an examination of the males and early stages may possibly prove ~ 7

it to be distinct.

The larvae (of the: typical form) were found in company with those of
A. annulipes Walk. and C. basicinctus Edw. associated with green algae in
running water in the Burnett River at Hidsvold. They were never found in any
other situation in this district. They differ from A. annulipes Walk. in the
following respects (Text-fig. 2 6b): The sublateral spine of the antenna is
definitely beyond the middle; the outer anterior clypeal hairs are single
(unplumed) and the posterior clypeal hairs are single, or sometimes bifurcate,
and extend well beyond the anterior margin of the head; the inner occipital hair
is single and the outer is trifid; the comb differs slightly in the arrangement of
the long spines, but is variable; in other respects, including the palmate hairs,
there is little or no distinction. Unfortunately the pupae were not preserved.

The biting habits and resting attitude of the adult females are, so far as my
experience goes, similar to those of A. annulipes Walk.

Distribution.—A. amictus Edw. has been recorded from North Queensland and
the Northern Territory. I have also taken it at Hidsvold, S. Queensland (April,
1924, bred from larvae, also biting after dusk) and there are specimens in the
Queensland Museum from the following localities in Queensland: Charleville,
June, 1899, C. J. Wild; EHidsvold, March, 1915, Bancroft; Brisbane, April, 1912,

Gb!
Nw 9?

38 * NOTES ON AUSTRALIAN MOSQUITOES, i.

H. Hacker; Cunnamulla, May, 1899, C. J. Wild. It would appear that in the
southern part of its range, which by the way is much more extensive than was
formerly thought, it is much less abundant and appears much later in the season
than further north.

ANOPHELES (ANOPHELES) BANCROFTI Giles.

There is no fresh information to add concerning this species. Adults and
the larvae are readily recognizable by the characters given in the keys. The
larva and the pupa have been described by Cooling (1921, 19240). The latter
stage is somewhat difficult to recognize, but is apparently similar to A. atratipes
Sk. from which it is to be separated by the markedly smaller paddles and probably
different chaetotaxy; further material is desirable. The male genitalia have been
described by Edwards (1924). Text-fig. 1 f was drawn from a male from the Sepik
River, Mandated Territory of New Guinea .(R. W. Cilento) which agreed in all
respects with typical females from the mainland and with Edwards’s description
of the hypopygium.

ANOPHELES (ANOPHELES) ATRATIPES Skuse.

A rather uncommon species which was hitherto known only from females.
Superficially it resembles A. bancrofti Giles, but is easily separable by the palpal
clothing and by the wing and leg markings. Its habits, too, are somewhat
similar in that it is a sylvan day-biting species which rests at an angle of about
80° with the surface. It is most prevalent in the coastal districts of New South
Wales and South Queensland in spring and early summer, apparently disappearing
completely later in the season when other Anophelines are most abundant. In the
Sydney district it occurs most frequently in the sandstone gullies and in the
vicinity of sluggish creeks running through the Pleistocene sand flats. At Dunwich,
Stradbroke Is., South Queensland, females were common in September, 1926, along
the course of a sluggish creek in which there were moderate numbers of the larvae.
A prolonged search was made among the vegetation, on the sheltered sides of
logs and tree trunks, and in similar likely situations, but not a single adult male
was seen. Neither males nor larvae have been found in any other locality, even
where females were not uncommon.

6. Palpi with the terminal segment swollen and with a patch of long hairs
ventrolaterally on the basal half of the terminal segment and extending on to the
apical part of the penultimate segment; antennae with long, dense, silky, brown
plumes; otherwise resembling the female in all essentials. The hypopygial
characters (Text-fig. 1 e) are in some respects intermediate between A. bancrofti
Giles and A. stigmaticus Sk., but show much closer affinities with the latter in
the single basal spine of the side-piece, in the pointed rather than lobed claspettes,
and in the replacement of the club by a row of four basal spines set on a lobe. It .
resembles A. bancrofti Giles in bearing three subequal spines on the claspette.
There are about two pairs each of longer medial and shorter lateral leaflets. The
allotype male (Dunwich, Stradbroke Is., South Queensland, bred from larvae,
10th Sept., 1926) is in the Macleay Museum, University of Sydney.

The larvae were taken in a small, sluggish, slightly muddy creek with a fine
silt bottom. The pH of a sample of the water was 7:5. (This determination was
kindly made for me by Dr. R. Hamlyn-Harris, City Entomologist, Brisbane, who
was also good enough to breed through a number of larvae brought back from
Dunwich.) Some aquatic vegetation was present, but not a great deal. Other

BY I. M. MACKERRAS. 39

mosquito larvae were entirely absent, although adults of several species were
common along the banks. The local distribution of the species in the Sydney
district suggests that it breeds in similar situations and also in pools in sandstone
creeks,

The larvae are darker than those of A. annulipes Walk., but do not afford any
naked eye characters of value for differentiating them. The antennae (Text-
fig. 2 d) are brown, notably stout and short, and bear apically two spines and a fine
trifid hair. The antennal hair is situated slightly basal to the middle, is fully
half the length of the antenna and is strongly branched. The head is very heavily
and uniformly pigmented and no trace of eyes could be made out in any of the
preparations. The anterior clypeal hairs resemble those of A. stigmaticus Sk., but
the posterior are very short and single. The frontal hairs are strongly plumed and
are subequal in length. The subantennal plume is normal. The inner occipital
‘hair is divided into four very fine branches. The outer lies well anterior to the
middle and is trifid. The thoracic and abdominal chaetotaxy is not remarkable
except for the entire absence or great reduction of the palmate hairs. These
structures may be represented by certain short hairs on a few of the abdominal
segments, but are certainly not present in the form seen in other Anopheline
larvae. The pecten consists in general of long spines alternating with two very
short ones. Grappling hooks are developed.

The affinities of the larva are with A. stigmaticus Sk., from which it is to be
separated by its larger size, much darker colour, much stronger antennal plume,
and by the absence of eyes or palmate hairs, a most remarkable and unexpected
feature of these larvae.

Text-figure 3.—Terminal segments of pupa of a A. annulipes Walk.; b A. stigmaticus
Sk.; ¢ A. atratipes Sk. Lateral view of trumpets of d A. annulipes Walk.; e A. stigmaticus
Sk.; f A. atratipes Sk.

The pupa appears to be closest to Cooling’s description of A. bancrofti Giles.
The skin is brown with strong lines of black pigment on the wing sheath along
the course of the veins and with transverse dark bars in a row aiong the antennal
sheath. The trumpet (Text-fig. 3 f) is very broad, triangular in shape, and is
deeply pigmented. This species is apparently separable from A. bancrofti Giles

40 NOTES ON AUSTRALIAN MOSQUITOES, i.

by its larger paddles and probably different abdominal chaetotaxy and trom all
other Australian species by the broad triangular trumpet, by the pigmentation of
the cephalothorax and by the chaetotaxy (Text-fig. 3 c).

It is interesting that this species should resemble A. bancrofti Giles fairly
closely in the general characters of the adults and in the pupa, but yet show
much closer affinities with A. stigmaticus Sk. on hypopygial and larval characters
which are undoubtedly more reliable.

ANOPHELES (ANOPHELES) STIGMATICUS Skuse.

This is a very rare, small brown species of which I have been fortunate in
obtaining a good series of both sexes and of the larvae and pupae at National
Park, New South Wales. They were only found in a patch of brush growing on
the Narrabeen series near Waterfall. Adults were taken by stirring them out
of the grass and bushes and at no time showed any inclination to bite. Their
resting attitude is almost horizontal and the palpi are carried closely appressed
to the proboscis so that they resemble when alive a small species of Culex much
more than an Anopheline.

Specimens were compared with the types of A. stigmaticus Sk. and found to
agree with them in every particular. The most striking features of this species
are the pale yellow basal four-fifths of the hind femora and the conspicuous dark
patch covering the forking of R and Rs. These characters are not mentioned
by Theobald in his description of A. corethroides Theo. and the stems of the
fork cells are distinctly longer than is indicated in his description. Skuse’s
description is complete and accurate except for the markings on the mesonotum’
which are correctly described by Theobald. A male in the Queensland Museum
from Burpengary, South Queensland (Bancroft) is from the type locality of
A. corethroides Theo. and agrees very well with his description. It agrees
completely in genitalic characters with typical A. stigmaticus Sk., but differs in
markings in that the dark patch on the wing is absent and the pale areas on the
hind femora are less extensive and creamy white rather than pale yellow. Of the
four other specimens from Queensland in the same collection, two are similar
to the male from Burpengary, one corresponds to the southern form, and one is
intermediate. These were all collected by Dr. Bancroft and are probably from
the vicinity of the type locality. The genitalia of both northern and southern
forms (Text-fig. 1 d is from a National Park specimen) agree with Edwards’s
(1924) description of the type of A. corethroides Theo. There is no doubt that
the synonymy given by Edwards is correct, but the typical corethroides form is
distinctly less well marked than typical stigmaticus.

The distribution is from South Queensland to National Park and the Blue
Mountains in New South Wales. The National Park specimens were taken in
April, 1925, and January, 1926.

The larvae were found associated with A. annulipes Walk. in small pools
containing filamentous algae in the bed of a small creek which ran through
the brush and had cut down to the Narrabeen sandstone. They are small, brown
in colour, and superficially resemble small specimens of A. annulipes Walk., naked
eye separation of the two being impossible. The antennae are creamy, darker at
the tip, and bear apically two spines and a single fine hair which is forked distally;
the antennal hair is situated at the junction of the basal and middle thirds of the
antennal length and is very short and finely branched. The head is uniformly
dark brown in colour and both compound and simple eyes are present. The inner

BY I. M. MACKERRAS. 41

anterior clypeal hairs are long, bare, and set close together; the outer anterior
clypeal hairs are a little more than half the length of the inner and are bare;
the posterior clypeal hairs are short and single or divided into two or three very
fine terminal branches. The frontal and subantennal hairs are normal. The inner
occipital hair is bifid and the outer lies well anterior to it and is trifid. The
chaetotaxy of the thorax and abdomen is not unusual. Palmate hairs occur on
segments 2-7 of the abdomen, are well developed and conspicuous, and each
consists of 20-25 leaflets which are long, slender, pointed, and not serrated. The
pecten is composed of regularly alternating long and short teeth. The ventral
subdorsal hairs bear the usual grappling hooks.

The pupal skin is a clear pale brown and the trumpet is not pigmented nor
are there any zones of pigmentation on the cephalo-thorax. This stage may
usually be separated from that of A. annulipes Walk. by its smaller size and paler
colour. The trumpet (Text-fig. 3 e) is longer and not so broad, and the abdominal
chaetotaxy (Text-fig. 3 6) is strikingly different. From A. atratipes Sk. it may be
distinguished by its smaller size, absence of pigmentation, much narrower trumpet,
and different chaetotaxy. The pupal stage lasted two to four days under the
conditions of the experiments.

References.

CoouIne, L. E., 1921.—On the larval and pupal stages of Myzorhynchus bancrofti Giles.
Proc. Roy. Soc. Q’land, 1921, pp. 166-173.

, 1924a—Seven common mosquitoes described for purposes of identification.
Cwlth. Dept. Hith. Serv. Publn. (Trop. Divn.) No. 1, 24 pp.

, 1924b.—Anopheline mosquitoes of North Australia and the tropical dependencies
of Australia. Preliminary notes. Appendix to Malaria in Australia. Cwlth. Dept.
Hlth. Serv. Publn. (Trop. Divn.) No. 3, pp. 97-141.

EDWARDS, F. W., 1924.—A synopsis of the adult mosquitoes of the Australasian Region.
Bull. Ent. Res., xiv, 4, pp. 351-401.

FERGUSON, EH. W., 1926.—Mosquitoes of New South Wales. 15th Rpt. Microbiol. Lab.
N.S.W. for Yr. 1924, pp. 187-191.

Hiuu, G. F., 1925.—The distribution of Anopheline mosquitoes in the Australian Region,
with notes on some Culicine species. Proc. Roy. Soc., Victoria, 1925, 1, pp. 61-77.
Money, R. A., 1926.—A case of malaria acquired in Sydney. Med. Jour. Aus., ii, 13th Yr.,

No. 9, 28th Aug., 1926, pp. 283-284.

THE INTERPRETATION OF THE RADIAL FIELD OF THE WING IN THE
NEMATOCEROUS DIPTERA, WITH SPECIAL REFERENCE
TO THE TIPULIDAE.

By CHarLes P. ALEXANDER, Ph.D."
(Communicated by Dr. E. W. Ferguson.)

(Ninety-two text-figures. )
[Read 25th May, 1927.]

Contents.
Historical.
The primitive condition of the radial field in the Diptera.
The two tendencies of specialization in the radial field in the Tipulidae.
A. Subfamily Limoniinae: Tribes Pediciini, Hexatomini and Eriopterini.
1. The cephalization of vein R,.
2. The capture of vein R, by R,,,.
3. The apparent radial crossvein in the Dicranotae.
B. Subfamilies Architipulinae, 'Tipulinae and Cylindrotominae; Subfamily
Limoniinae: Tribes Lechriini and Limoniini.
1. The serial nature of vein R,.
Phylogenetic considerations.
Summary.
References.

HISTORICAL.

The study of the venation of the Diptera during the past century has brought
about many successive and progressive modifications. The early interpretations
of Schiner (1862, pl. 2), Loew (1862) and Osten Sacken (1869) were materially
improved by Williston (1908, pp. 37-43) and other subsequent workers. The
proposal of the Comstock-Needham system of nomenclature and its application
to the order (1899) furnished the most important means for a true knowledge and
appreciation of the homologies of the veins. Still more recent work by Tillyard
(1919) has served to strengthen our interpretation of the medial and cubital fields
of the wing.

In the course of his studies on the Tipulidae, the present writer has long felt
dissatisfied with the hitherto adopted interpretation of the radial field of the wing
in Diptera (Needham, 1908; Comstock, 1918, 1924; Tillyard, 1919; MacGillivray,
1923). At first sight this interpretation would suggest a double dichotomy of the
branches of the radial sector, as in the hypothetical type of insects, and this is
indeed true in the two most generalized families of living Diptera (Tanyderidae
and Psychodidae). In all higher Diptera, however, one or more branches of this
dichotomy have been lost and this has been interpreted as having been brought
about by a fusion to the margin of either the upper fork (Ptychopteridae;

1Contribution from the Department of Entomology, Massachusetts Agricultural
College, Amherst, Massachusetts, U.S.A.

\

BY CHARLES P. ALEXANDER. 43

Brachycera) or the lower fork (Nematocera) of the primitive dichotomous sector.
It has long seemed very strange that in the Diptera we should start with two
extremely primitive families (Tanyderidae and Psychodidae) that have no
indication of a radial crossvein and then suddenly gain such a vein in the Tipulidae
and a few more or less closely allied groups, following which the structure has
been completely lost throughout the order. In the present paper, the writer has
attempted to demonstrate that the true radial crossvein (r of Comstock (1918,
1924), Needham (1908); ir, of Tillyard) has never been developed as a transverse
element in the Diptera, as shown by all of the above students, and should be
omitted as such from any archetype of the order (Tillyard, 1919, fig. 69). The
vein lies as a longitudinal element in a serial alignment, distinct as a separate unit
only in the subfamilies Architipulinae, Tipulinae and Cylindrotominae, and two
tribes of the Limoniinae, the Lechriini and some Limoniini. In two earlier papers,
the writer had briefly outlined these various interpretations in the subfamily
Cylindrotominae (1919, p. 863) and in the Limoniine tribe Pediciini (1918).

Edwards (1926, p. 126) felt inclined to doubt the possibility of two radically
different types of radius having been developed within the family Tipulidae. It
was for the purpose of attempting to decide this question, which has long been a
disputed point in the consideration of the group, that the present writer began a
critical comparative survey of the various genera and species, more than 98%
of the genera and subgenera, and over 3,000 species of Tipulidae having been avail-
able for the purpose. It was soon apparent that two distinct groups were involved,
one, represented by the Tipulinae, Cylindrotominae, and the tribes Lechriini and
Limoniini, having r present as a longitudinal element and forming part of a serial
radial vein and with R, and R, fused to the wing margin; and, secondly, the
remaining tribes of the Limoniinae (Pediciini, Hexatomini, Hriopterini, and the
aberrant Styringomyini) in which no radial crossvein had been developed, where
R, had swung cephalad and fused backward from the wing-margin with R,, where
R, had been taken from the primitive dichotomous posterior fork of the sector
and become more intimately connected with the anterior branch, R,,,, leaving the
posterior branch of the radial field to consist of R, alone. This rather involved
and somewhat mystifying condition of affairs is discussed in some detail in the
present paper. A special effort has been made to choose representatives of
Australian genera where these showed the points involved. After this critical
review of the known Tipulidae, the writer cannot feel but that the main principles
as given at this time are correct.

THE PRIMITIVE CONDITION OF THE RADIAL FirLpD IN THE DIPTERA.

The condition of the radial field in the two recent families Tanyderidae and
Psychodidae is so like that of the archetype of the Diptera that we must consider
them as representing the two most generalized living groups. This contention is
beautifully upheld by the recent critical work on the body morphology and
phylogeny of the group by Crampton (1924-1926) and Edwards (1926). There
can be no doubt but that the two families are closely allied, a view that was
apparently first suggested by Lameere (1906). In these two groups there are five
branches of radius attaining the wing-margin and no indication of a radial cross-
vein. It should be noted, however, that in the three families of Diptera most
closely allied (Ptychopteridae, Trichoceridae and Tipulidae) the number of
branches of the sector has already undergone a reduction and in representatives
of all of them the so-called radial crossvein (7 of previous workers) is found.

44 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

In the phylogeny of the Diptera (Crampton, 1924-1926; Edwards, 1926) it is now
believed that the Ptychopteridae are closely allied to the Tanyderidae and that
they either ended as a blind stem, or, possibly, gave rise to certain groups of the
Nematocerous Diptera, and through them, the Brachycera. The Trichoceridae
appear to lead directly toward the Anisopodidae, from whence the other groups
of the Nematocera seem to have been derived. Since, therefore, the only groups
of Diptera that are more primitive than these three are the Tanyderidae and
Psychodidae, it follows that whatever interpretation is accepted for the
Ptychopteridae, Trichoceridae and Tipulidae must be accepted also for all higher
Diptera. It is for this reason that the correctness or incorrectness of the accom-
panying data and interpretations assumes an importance very much greater than
would be the case if merely the single family Tipulidae were involved.

Ture Two TENDENCIES OF SPECIALIZATION IN THE RADIAL FIELD IN THE TIPULIDAE.

As was briefly indicated in the historical introduction of this paper, there
appear to have been two very distinct tendencies in the Tipulidae that are so
profound that both are shown by the oldest known Tipulidae and related groups
(Upper Liassic).

These two lines of specialization are as follows:

A. In the subfamily Limoniinae, the three tribes Pediciini, Hexatomini and
Eriopterini (together with the aberrant tribe Styringomyini, which apparently
must be considered with the Eriopterini in any discussion of the groups) have
evidently been derived through some primitive Pediciine ancestor from a type
that was not unlike our recent Tanyderidae. The upper branch of the anterior
fork of the primitive sector has swung cephalad and become fused backward from
the margin with vein R,; in all but a few generalized members of these three
tribes the dichotomous nature of the posterior fork of the sector has been lost
by the capture of the anterior branch (R,) by the stem of the upper fork
(R,,,), leaving R, alone as the posterior branch of the radial field.

B. In the subfamilies Architipulinae, Tipulinae, Cylindrotominae; and the
tribes Lechriini and Limoniini of the Limoniinae, a radial crossvein is present,
occupying a longitudinal or oblique position, and in all higher types becoming
part of a serial vein composed of R, +7 +R, The distal sections of R, and R, are
lost by atrophy except in the more generalized groups. As in the preceding
section A, the apparent radial crossvein is really the basal section or deflection of
R,, in the higher groups of the Limoniini this deflection being lost by atrophy.
The posterior fork of the primitively dichotomous sector is interpreted as being
lost by a fusion of the two elements, R, and R, to the wing-margin. There is no
direct evidence to support this interpretation, however, and it must be held as
somewhat tentative (see discussion in concluding paragraph of the
Cylindrotominae).

A. Subfamily Limoniinae: Tribes Pediciini, Hexatomini and Eriopterini.
1. The cephalization of vein R,.

In the primitive families of Diptera, the Tanyderidae and Psychodidae, vein
R, is a full length vein that lies parallel to veins R, and R, and attains the wing-
margin as a separate entity. In all other groups of Diptera, vein R, is either
connected with R, by the longitudinal crossvein r (as discussed in detail under
caption B) or has swung cephalad and forms a fusion with R, backward from
the margin.

BY CHARLES P. ALEXANDER. 45

As a preliminary to the data that are presented in this paper, the family
Tanyderidae should be briefly considered since it is obviously closest to the
ancestral type from which the Pediciini and higher Limoniinae have been evolved.
The family includes the following genera: Macrochile Lw., fossil, Lower Oligocene
(Crampton, 19260); Protoplasa O.S., 3 Nearctic, 1 Palaearctic species;
Péringueyomyina Alex. (barnardi Alex.), Ethiopian; Tanyderus Phil. (pictus
Phil.), Neotropical; Radinoderus Handl., 7 species, chiefly Australian, 1 Neo-
tropical; Mischoderus Handl., 4 or 5 valid species, all from New Zealand; Neoderus
Alex. (patagonicus Alex.), Patagonia; and Nothoderus Alex. (australiensis Alex.)
of Tasmania. Nothoderus australiensis Alex. was discussed and figured by

M, M, 4 :

Fig. 2 2nd A Ista Cu,

Text-figure 1. Wing of Nothoderus australiensis Alex. (Tanyderidae).

Text-figure 2. Wing of Tricyphona protea Alex. (Pediciini).

A = anal veins; C = costa; Cu = cubitus; h = humeral crossvein; M = media;
m = medial crossvein; m-cu = medial-cubital crossvein; R = radius; r-m = radial-medial
crossvein; Rs = radial sector; Sc = subcosta; s = supernumerary crossvein in cell Ry,

MacGillivray (1923, p. 322, fig. 45) as being “the most generalized dipterous wing
that I have seen’’, despite the fact that the free tip of Sc., a remarkable feature
of the genus, is omitted from his figure. This primitive fly (Text-fig. 1) shows the
arrangement of the branches of radius in the Tanyderidae. The supernumerary
crossvein (s) in cell R, is a feature that is very characteristic of the members of
this family and likewise of the Pediciini.

The first realization of the cephalization of vein R. in the lower Diptera came
to the writer with the discovery of a remarkable crane-fly, Tricyphona protea Alex.
(Text-fig. 2) where the venation is very like a Tanyderid except that vein R, has
swung cephalad and fused with R, for a short distance back from the wing-margin.

46 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

The study of this fly and a comparison with the then known members of the
tribe Pediciini resulted in a short paper by the writer (Alexander, 1918) wherein
the radial field in this tribe was interpreted about as is done in the present treat-
ment. This interpretation for the Pediciini was adopted by Edwards (1921) and
the writer but no attempt was later made to homologize the venation with that
of the other tribes in the Limoniinae and Tipulinae and, from them, the phylo-
genetically higher families of Diptera. A second species of Tricyphona (formosana
Alex., Text-fig. 3) was later discovered in which the venation of this field is about
intermediate between protea and the normal condition in the tribe (7. novae-
zelandiae Alex., Text-fig. 4) where this element (the basal section of R,) is short

Sc, R, R

2nd A My

Fig. 3 Fig. 4

Text-figure 3. Wing of Tricyphona formosana Alex.
Text-figure 4. Wing of JT. novae-zelandiae Alex.

and transverse in position. It is this basal section or deflection of R. that has
almost universally been interpreted as being 7 in the groups of Diptera where it
has been retained. In a few of his recent papers, the present writer has indicated
his opinion that 7 as a transverse element did not exist in the Tipulidae or in the
Diptera.

2. The Capture of Vein R, by Rz..

We ‘now come to one of the most curious features in the problem. In the
primitive Diptera (Tanyderidae, Text-fig. 1; Tricyphona protea Alex., Text-fig. 2),
as elsewhere in generalized groups of the higher orders of insects, the radial sector
is typically dichotomously twice forked. In the Tipulidae this dichotomous type
has been preserved only in a few scattered groups, chiefly in the Pediciini (Text-
figs. 8, 9), and in some Eriopterini (Molophilus, Text-fig. 39), the upper fork in
all of these types being eliminated by the fusion of the upper branch with R,, as
just outlined. The lower dichotomy in these groups retains its primitive condition
of a very deep fork and correspondingly shortened petiole. However, in many
members of the tribe under consideration, the Pediciini, as, for example, in the
genus Rhaphidolabis (Text-figs. 5-6) we find a bewildering condition where the
dichotomy is lost by R, becoming more intimately attached to the upper fork than
to the lower. Similarly in the genera most closely allied to Molophilus, as
Erioptera, we find an entirely comparable condition. This apparently irreconcilable
condition is really brought about in a very simple manner by a slight shifting of
the relative positions of the veins at the end of the sector. This condition was
discussed at some length by Needham in his classic study of the venation of the
Tipulidae (1908, pp. 225-226, fig. 14) and has been investigated more superficially by
subsequent workers on the family. For the purposes of illustration of the principle,

BY CHARLES P. ALEXANDER. 47

species of the Pediciine genus Rhaphidolabis O.S. in the Nearctic fauna have been
selected (Text-figs. 5-9). In species like R. cayuga Alex. (Text-fig. 9, 9a) and
R. stigma Alex. (Text-fig. 8), the primitive condition of the lower fork of the
sector is found, cell R, being broadly sessile, cell R, petiolate. In &. rubescens
Alex. (Text-fig. 7, 7a) the petiole of cell R, has shortened so that both cells R,
and R, appear sessile, vein R, being in direct alignment with the sector. From this
point on, vein R, moves progressively upward on to the stem of the upper fork,
R,,,, In R. major Alex. (Text-fig. 6, 6a) this migration is very slight so that cell

acl
R,
R,
Fig. 5
co
SS Bie
Fig, 9
Fig. 6 , °
Rs a ‘ da

Se 6a
Rs-
ae
aS Rs =. 243 P
Peas

Fig, 7 R

Text-figure 5. Wing of Rhaphidolabis polymeroides Alex.

Text-figure 6. Wing of R. major Alex.

Text-figure 7. Wing of R. rubescens Alex.

Text-figure 8. Wing of R. stigma Alex.

Text-figure 9. Wing of R. cayuga Alex. Subfigures, ia, 6a, 7a and 9a, showing
details of the forking of the sector in the above species.

R, would be described as being short or very short petiolate, the fused vein
R2,3,, being this petiole. In still other species, illustrated by R. polymeroides Alex.
(Text-fig. 5, 5a), the capture of vein R, by R,,, has been more complete and the
petiole of cell R, is conspicuous. Parallel cases are to be found in Tricyphona
Zett., Nipponomyia Alex. (Text-figs. 14-16) and elsewhere in this generalized tribe.
The end result of this shifting is the basal fusion of R.,,, with R,; which becomes
very extensive in some groups of the family. As the fusion progresses, cell R;,

48 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

becomes more shallow until, in cases discussed under the Hexatomini and
Eriopterini, it is entirely lost by the fusion to the wing margin of veins R, and
R,. After the capture of vein R, in the manner described, the single remaining vein
of the radial field is R,.

3. The Apparent Radial Crossvein in the Dicranotae.

A few Pediciini (all species of Dicranota, Amalopina dicranotoides Alex. and
A. sibiriensis Alex., Polyangaeus) possess a crossvein in cell R,, which, in an
earlier paper by the writer (1918, p. 205) was interpreted as being the true r.
In the light of the evidence now available that .. in the Diptera, in the relatively
few groups where it occurs (as described under caption B, later) always occupies
a longitudinal or oblique position and becomes one of the series of elements
making up the serial radial vein, it becomes necessary to examine the case in the

Fig. 10

Fig. Il

Text-figure 10. Wing of Dicranota bimaculata Schumm.
Text-figure 11. Wing Heterangaeus gloriosus Alex.
R = radius; Se = subcosta; s = supernumerary crossveins.

Dicranotae more critically. It should be noted first that this tribe shows a
plasticity in the number and position of crossveins, both supernumerary and
adventitious, quite unparalleled elsewhere in the Tipulidae, though virtually
duplicated in the Tanyderidae. The genus Heterangaeus Alex. (Text-fig. 11) shows
how abundant these crossveins may be in a single genus and species and there |
is no reason to believe but that the crossvein in cell R, in the cases above
outlined, is merely another instance of a supernumerary crossvein, in this case
occupying a cell where its position and interpretation becomes far more critical
than in the other instances. The plasticity of the character is shown by the fact
that Amalopina dicranotoides and A. sibiriensis are the only ones of the genus (of
six species) that possess this supernumerary vein. The genotype, elegantula

BY CHARLES P. ALEXANDER. 49

Brun., lacks this vein but has another in cell R, Dicranota (Text-fig. 10), in
which this character is present, is very closely allied to Rhaphidolabis O.S.
(Text-figs. 5-9) in which the crossvein is lost, and the presence or absence of this
element constitutes the only criterion for the separation of the two groups.
Taking into consideration the unusual degree of occurrence of supernumerary
and adventitious veins (Johnson, 1901) in members of this tribe, there can he
no serious objection to considering the vein in the Dicranotae as being merely
an additional supernumerary element stranded in cell R,.

The writer now wishes to take up the three tribes in this group and discuss
the more interesting tendencies. j

Pediciini.—As was indicated, the primitive arrangement of the veins at the end
of the sector is found in many groups of the tribe. In Pedicia Latr., Ornithodes Coq.,
most Tricyphona Zett. (Text-figs. 2-4, 12, 13), as well as in Rhaphidolabis O.S.
(Text-figs. 8-9) the dichotomous arrangement of the veins still obtains. The petiole
of the posterior fork (R,,;) is usually short, reaching its maximum length in species

: bs | ? Roneeae R,..,
sree 70)
R,
Fig. 13 a!

Text-figure 12. Wing of Tricyphona brevifurcata Alex.

Text-figure 13. Wing of T. rainieria Alex.

Text-figure 14. Wing of Nipponomyia kuwanai Alex.

Text-figure 15. The same, showing modification of veins at end of sector.
Text-figure 16. Nipponomyia symphyletes Alex.

such as 7. brevifurcata Alex. (Text-fig. 12). The remarkably primitive condition
found in 7. protea Alex. (Text-fig. 2) has been discussed elsewhere in this paper.
No species is definitely known where R, ends beyond the level of R, and connects
directly with costa. In 7. rainieria Alex. (Text-fig. 13) the deflection of R, is at
the extreme outer end of vein R,, as is also the case in Polyangaeus maculatus
Doane. Doane even figures his type example of this latter species as having this
element connected with costa but no mention is made of this very unusual
condition in the text and it is presumably erroneous.

The manner in which the dichotomous posterior fork is lost has been discussed
earlier in the case of Rhaphidolabis (Text-figs. 5-9). A closely comparable case is
found in the species of the genus Nipponomyia Alex., where the various modifica-
tions are found at times in different individuals of the same species. Thus
N. kuwanai Alex. may have cell R, short-petiolate (Text-fig. 14) or sessile (Text-

F

50 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

fig. 15). N. symphyletes Alex. (Text-fig. 16), N. trispinosa Alex., N. novem-
punctata S.-W., and N. sumatrana de Meij., have R, mounted half-way up the
perpendicular basal portion of R,,, so that cell R,; has become petiolate, cell R,
broadly sessile. In the genus Jricyphona we find a great group of species in which
the primitive condition is still retained but in still others (as inconstans O.5S.,
constans Doane, and many of the European species, as gmundensis Hgg., littoralis
Meig., occulta Meig., opaca Hgg., schineri Kol., tipulina Egg., and others) R, has
been captured by R..3 in the manner previously described. It should be noted
here that occasional specimens of inconstans are found that show a reversion to the
primitive condition of the genus (Osten Sacken, 1869, p. 267; Johnson, 1901,
p. 307). Specimens in the related genus Pedicia show a similar range in this field
of the wing. As was indicated earlier in this paper, supernumerary crossveins are
commonly developed in this tribe, occurring in cells R, R, R, and M in
Polyangaeus and:in cells R,, Ru, M, and M in Heterangaeus.

In the subtribe Adelphomyaria, including only the genus Adelphomyia Beregr.,
a connecting element with the following tribe, the Hexatomini, is found. The
adult flies are very Hexatomine in structure but the larvae exhibit some curious
Pediciine characters. The venation of the radial field is not different from
Limnophila, discussed later, except that the basal section of R, is often weakly
developed and tends to be lost by atrophy.

Hexatomini. This tribe is apparently naturally derivable from the Pediciini
through the Adelphomyaria on the one hand, and the lowermost subtribe, the
Ularia, on the other. Within the tribe there is found a remarkable range of
usually large to medium-sized species that are arranged in a variety of subtribes.
The number of these latter sub-divisions will probably be increased when certain
doubtful genera become better known and new types are discovered. The most
specialized groups appear to fall. in the Gynoplistiaria, the Limnophilaria, the
Hexatomaria and the Elephantomyaria, some of which lead rather gradually
toward the Eriopterini. The chief modifications and tendencies in the radial field
are discussed below.

Ularia. Includes only the genus Ula Haliday. The venation is not different
from some species of Tricyphona, as inconstans O.S8.

Epiphragmaria. Includes a number of chiefly Antipodal genera, as Wpiphragma
O.S., Polymoria Phil., Limnophilella Alex., Acantholimnophila Alex., Rhamphophila
Edw., Tinemyia Hutt., Phyllolabis O.S., Horistomyia Alex., Heterolimnophila Alex.,
Polymera Wied., and others. The venation shows no modifications of the radial
field over that of Gynoplistia, described later.

In Polymera, the position of the basal section of R, differs greatly in the
various species, in some, as inornata Alex., the last section of R, being equal to
R,., while in other species the section of R. is far out toward the tip of R,.

Pseudolimnophilaria. Includes only the genus Pseudolimnophila Alex., as
known, but other groups will probably be added to the subtribe when better known.
The genus included is not different in venation from Limnophila.

Dactylolabaria. Includes only the genus Dactylolabis O.S., so far as known.
As before, the venation is normal for the Hexatomini and requires no comment.

Gynoplistiaria. Includes only the genus Gynoplistia Westw., with four
subgeneric groups, Paralimnophila Alex., Gynoplistia, Cerozodia Westw., and
Ctedonia Phil. The group is tremendously developed in Australia and New
Zealand. The figure of G. bella Walk. (Text-fig. 17) may be taken as typical not
only of the subtribe but for the entire tribe Hexatomini. R.,;,, is almost always

BY CHARLES P. ALEXANDER. 51

very short to entirely obliterated (as in G. dimidiata Alex., G. harrisi Alex
G. hiemalis Alex., G. nigronitida Alex.), cell R, thus varying from short-petiolate
to sessile; R,,, elongate, usually perpendicular or subperpendicular at origin,
sometimes with a short spur or weak crossvein at the angle; basal section of
R, short, the fused R,,. generally short to of moderate length.

Limnophilaria. This subtribe as now constituted includes a considerable
range of genera, some of which are discordant and will later be removed. A few
of the groups require special comment. Psaronius End. shows a venational range

Fig, 17

M,
2nd A Ist ‘A Cu,

Text-figure 17. Wing of Gynoplistia bella Walk.

of the radial field that closely parallels that of the Hriopterine genera
Paragymnastes and Gymnastes. Rs is long and lies close to R, in alignment with
R».3,, and its continuation, R,; even in the most generalized species, as obscurus
Fabr. (Text-fig. 18), R.,, is relatively short and R, weak although long and lying
parallel to the fused R,,,.. In trianguliferus Alex. (Text-fig. 19) the tip of R, has
swung basad and is fused with the tip of R,,., cell R, being elongate-triangular.

Ri

Ri4243
=}
Fig. 18 : Fig. 20
ae
R, a
Fig. 19 Fig. 21 |

Text-figure 18. Wing of Psaronius obscurus Fabr.
Text-figure 19. Wing of P. trianguliferus Alex.
Text-figure 20. Wing of P. obliteratus Alex.
Text-figure 21. Wing of P. abnormis Alex.

In obliteratus Alex. (Text-fig. 20) the fusion backward from the tip has proceeded
so far that cell R. is reduced to a tiny triangle. In species such as abnormis Alex.
(Text-fig. 21), brevitibia Alex., and mancus Alex., the tendency is carried to its
logical conclusion and cell R, is entirely obliterated.

52 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.
The genus Jschnothrix Bigot is of importance as indicating the manner in

which the aberrant genus Atarba O.S. has been derived. In J. australasiae Skuse
(Text-fig. 22) R...,, is very long, the branches R,; and R, short but separate, cell

Rene 3

Text-figure 22. Wing of Ischnothrix australasiae Skuse.
Text-figure 23. Wing of J. connexa Alex.
Text-figure 24. Wing of Atarba picticornis O.S.

R,; being small. In still other species, as I. connexa Alex. (Text-fig. 23) a secondary
modification is produced by R, having swung basad and fusing backward from
the margin with R,,., presumably for the purpose of strengthening this field of the
wing. Similar cases are found in the Eriopterine genus Gonomyia, subgenus

Fig. 26 UE

Fig. 29 :

Text-figure 25. Wing of Hriocera velveta Doane.
Text-figure 26. Wing of H. wilsoni O.S.
Text-figure 27. Wing of H#. gibbosa Doane.
Text-figure 28 Wing of H. fultonensis Alex.

Ls)
©

Text-figure Wing of Hexatoma megacera O.5.

ol
es)

BY CHARLES P. ALEXANDER.

Ptilostena Bergr. In Atarba (picticornis O.S., Text-fig. 24) veins R, and R, have
fused to the wing-margin or, possibly, one of the branches atrophied shortly before
the margin.

Hexatomaria. This very extensive and important subtribe includes four very
closely allied genera, Hriocera Macq., Penthoptera Schin., Hexratoma Latr., and
Cladolipes Lw. Members of this subtribe are chosen to illustrate the manner in
which veins R.,, and R, fuse to the wing-margin in certain groups of Hexatomini
and Eriopterini. The figures show a series of Nearctic species of Hriocera, with
one species of Hexatoma. In E. velveta Doane (Text-fig. 25) cell R; is still very
deep; in H. wilsoni O.S. (Text-fig. 26) the cell is a little shorter in proportion to its
petiole; EH. gibbosa Doane (Text-fig. 27) shows the fusion as having progressed to
opposite the point of departure of the basal section of R.; H. fultonensis Alex.
(Text-fig. 28) shows the fusion of R, and R, as having continued to some distance
beyond R.; Hexatoma megacera O.S. (Text-fig. 29) has the veins fused almost to
the margin. The condition results in a complete fusion in the Huropean Cladolipes
simplex Lw. The deepest cell in this subtribe known to the writer is in EL.
subrectangularis Alex., where R,,;,, is about as long as 7-m.

Elephantomyaria. The very aberrant genus Hlephantomyia O.S. (westwoodi
O.S., Text-fig. 30) is placed in the Hexatomini with some question although the
reference seems to be correct. The genus as interpreted shows veins R, and R,

Re Rass
RK R,
Fig. 30 Fig. 3]

Text-figure 30. Wing of Hlephantomyia westwoodi O.S.
Text-figure 31. Wing of H. major Alex.

fused to the wing-margin and the basal section of R. entirely atrophied. In the
subgenus Hlephantomyodes Alex., of the Indo-Australian Region (major Alex.,
Text-fig. 31), R.,.,, at origin is perpendicular to subperpendicular, the distal section
being closely approximated to R,.

In the tribe Hexatomini, the basal section of R. is lost by atrophy in many
genera, as Horistomyia leucophaea Skuse, Phyllolabis O.S., some Zelandomyia Alex.,
Harrisomyia Alex., Ischnothrix Big., Hlephantomyia O.S., and others. A few
groups show a supernumerary crossvein in cell R,, as Ctenolimnophila Alex.
(decisa Alex.); the subgenus Calolimnophila Alex., of Pseudolimnophila Alex.; the
subgenus Dicranophragma O.S. of Limnophila Macq.: Elephantomyia super-
numeraria Alex., and others. In Ctenolimnophila bivena Alex., there is an
additional supernumerary crossvein in cell Ive

Hriopterini. The great tribe Eriopterini has evidently been derived from
the Hexatomini through certain of its subtribes, as the Claduraria and Conosiaria
which have retained numerous Hexatomine features and have been referred to as
the ‘“Hexatomoid Eriopterini.” The most generalized subtribes would thus be those
just mentioned, leading, through the Gonomyaria to the so-called ‘“pot-bellied”’
Eriopterini, the typical Eriopteraria. -In this vast assemblage of genera and sub-
genera several specializations and modifications from the normal type are found
that are very puzzling and difficult of explanation. The greatest trouble comes in

54 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

three, chiefly Ethiopian, genera, Ceratolimnobia, Xipholimnobia and Xenolimnobia,
_ which, as discussed later, may well pertain to the Lechriini in the following group
of subfamilies. More material will be needed to determine finally the exact
systematic position of many of the crane-fly genera and a critical study of all the
stages and available structures must be made. Very many of the larger and more
widely distributed genera are vastly ancient and there has been almost unlimited
time during which the various modifications and tendencies have been evolved.
It is very certain that more subtribes will be formed when certain groups of
exotic Eriopterini become better known. In many of the higher members of the
tribe, the basal section of R, has atrophied, leaving no hint as to its exact former
position. When present, this element shows a great range in position, in some, as
Molophilus (Text-fig. 39) and Hrioptera (Text-fig. 40) being retreated far from
the tip of R,, the fusion of R, and R, thus being very extensive; in still others
the element is far out toward the end of vein R,. When the element has atrophied,
as in Gonomyia and many other groups, all that can be affirmed is that the distal
section of R, has fused with R, and the vein at the margin must be called R,,..

In the present paper, only a few of the more conspicuous types and certain
modifications can be discussed.

Claduraria. This subtribe includes Cladura O.S., Neocladura Alex., Crypteria
Bergr., Pterochionea Alex., Neolimnophila Alex., and Chionea Dalm., a group of
genera very characteristic of the northern Holarctic Region, most of them appear-
ing as adults in autumn and winter. The venation shows no modifications from
the normal type of the Hexatomini and is not further discussed.

Conosiaria. Includes Conosia v.d.W. and Clydonodozus End., two closely
allied Palaeotropical genera in which the maxillary palpi are greatly reduced.
The more generalized genus, Clydonodozus, has the basal section of R, transverse
and not greatly different from the condition in the Hexatomini (angustifasciatus
Alex., Text-fig. 32). The more specialized genus, Conosia (irrorata Wied., Text-

R,

Fig. 32 i net 4

Text-figure 32. Wing of Clydonodozus angustifasciatus Alex.
Text-figure 33. Wing of Conosia irrorata Wied.

fig. 33) has the basal section of R, reflexed back toward the wing-base so that the
proximal end of cell R, is acutely pointed. It should be noted that the shifting
of veins at the end of Rs, as already discussed in some detail for the Pediciini, is
found in these two genera and even in different specimens of the single genus
Conosia.

Gonomyaria. An extensive subtribe, interpreted as including, among others,
Gonomyia Meig., Gnophomyia O.S., Teucholabis O.S., Trentepohlia Big.,
Paragymnastes Alex., Gymnastes Brun., Campbellomyia Alex., Dasymallomyia
Brun., Lipsothriz Lw., Aphrophila Edw., Astelobia Edw., Austrolimnobia Alex.,
Sigmatomera O.8., Lecteria O.S., Platylimnobia Alex., and more doubtfully,
Ceratolimnobia Alex., Xipholimnobia Alex. and Xenolimnobia Alex.

BY CHARLES P. ALEXANDER. 55

The following groups should be more critically examined to determine their
strict position in subtribes; (1) Trentepohlia; (2) Sigmatomera, Astelobia,
Austrolimnobia; (3) Lecteria (and its exact relationships with the Hexatomine
genus Psaronius); (4) Paragymnastes and Gymnastes; (5) Aphrophila; and (6)
Ceratolimnobia, Xipholimnobia and Xenolimnobia. The genera as listed above
are grouped chiefly on thoracic characters and, as so united, unquestionably include
some discordant elements.

Most of the groups require no special explanation to homologize their venation
with the groups previously discussed. In Trentepohlia and Astelobia, the basal
section of R, is reflexed strongly basad, as in Conosia, just described. The allied
genus Austrolimnobia has the element transverse and normal in position. In
Paragymnastes (fascipennis Thoms., Text-fig. 34) and Gymnastes Brun.
(ornatipennis de Meij., Text-fig. 35, shirakii Alex., Text-fig. 36) we find the loss of
the free distal section of R, by gradual atrophy and probable fusion back from

R,
Sc Sc, Riss R

Fig. 36

Text-figure 34. Wing of Paragymnastes fascipennis Thoms.
Text-figure 35. Wing of Gymnastes ornatipennis de Meij.
Text-figure 36. Wing of G. shirakii Alex.

Text-figure 37. Wing of Xipholimnobia terebrina Alex.
Text-figure 38. Wing of Xenolimnobia camerounensis Alex.

the tip of R,,., much as previously described under the Hexatomine genus
Psaronius. In Gonomyia and Teucholabis we find a progressive increase in the
degree of fusion of veins R; and R,, in exactly the same manner as discussed under
Hriocera (Text-figs. 25-29) in the preceding tribe. This culminates in the subgenera
Lipophieps Bergr. and Teucholabis, respectively, cell R, being obliterated, producing
a venation that is very similar to that found in the Limoniini in the next section.
In Teucholabis, r-m is sometimes obliterated by the fusion of vein R; on M,,..

In Ceratolimnobia and allied genera, as mentioned above, a much more
difficult and serious problem is found. In Ceratolimnobia and Xipholimnobia
(terebrina Alex., Text-fig. 37) the appearance is much as in TJrichoneura, that has
been placed in the Lechriini of the following section, except that the critical

56 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

distal section of R, is not preserved in any material that has been discovered
and the apparent homology may be apparent only. In Xenolimnobia
(camerounensis Alex., Text-fig. 38) the condition is reversed, the apparent tip of
R, being preserved but the entire distal section of R. atrophied. It is greatly
hoped that additional and possibly annectant forms in this group will be
discovered.

Eriopteraria. This subtribe is restricted to the so-called “pot-bellied”
Eriopterini (Crampton, 1925, p. 199) and includes among others, the following
genera: LHrioptera Meig., Empeda O.S., Psiloconopa Zett., Cryptolabis O.S.,
Dasymolophilus Goetg., Tasiocera Skuse, Molophilus Curt., Ormosia Rond.,
Amphineurus Skuse, Trimicra O.8., Helobia St. Farg., Podoneura Berer.,
Empedomorpha Alex., and Rhabdomastix Skuse. The species of Rhabdomastiz
bear a noteworthy resemblance to Gonomyia, with which genus they were long
confused in the European fauna. The genus entirely parallels Gonomyia in that
some few generalized species have retained the basal section of R, while in the
majority it is entirely atrophied. In all of the species of the genus yet made
known, cell R,; is present though often very small.

In Molophilus (froggatti Skuse, Text-fig. 39) we find the same generalized
dichotomy of the sector that has been considered under the Pediciini. An inter-
mediate condition is found in Dasymolophilus and, especially, in several New
Zealand species of Amphineurus where cell R, ranges from barely sessile through
other species where the cell is short-petiolate, to Er ioptera (lutea Meig., Text-fiz.

Rizr

Fig. 4I

Text-figure 39. Wing of Molophilus froggatti Skuse.
Text-figure 40. Wing of Hrioptera lutea Meig.

Text-figure 41. Wing of Empedomorpha empedoides Alex., ¢.
Text-figure 42. Wing of Styringomyia terrae-reginae Alex.

40) where the cell is distinctly petiolate. A curious modification of the veins of
the radial field in the male of Empedomorpha (empedoides Alex., Text-fig. 41) is
found where the basal section of R, is very long, the other elements being reduced
and crowded. The female of the same species is a very ordinary fly.
Styringomyini. The single very isolated genus Styringomyia Lw. (terrae-
reginae Alex., Text-fig. 42) is placed here. The venation of the radial field some-

BY CHARLES P. ALEXANDER. 57

what suggests Ceratocheilus, as discussed under the following section, R.,, being
very short and lying obliquely, far distad of R, Any discussion as to how this
remarkable venation might have been evolved would be theoretical only.

B. Subfamilies Architipulinae, Tipulinae and Cylindrotominae; Subfamily
Limoniinae: Tribes Lechriini and Limoniini.

1. The serial nature of vein R,.

The term serial vein was proposed by Comstock (1918, p. 69) for a compound
vein composed of sections of two or more veins joined end to end with no indica-
tion of the point of union. In the subfamilies and tribes considered under this
caption, the vein that has hitherto been called R, (Needham, 1908; Comstock,
1918) is, in reality, such a serial vein, being composed of three distinct elements,
R,, 7, and the distal section of R,. At the margin the vein should be labelled as
being R, alone (except in the cases noted below), and the ¢ell beneath this distal
section of R, should be called R. instead of cell 2nd R, as heretofore. In the
Tipulinae, Cylindrotominae, Lechriini and the more generalized Limoniini, the
three elements constituting the distal end of this serial vein still preserve much
of their separate identity by a more or less zig-zag arrangement, and, in many
cases, by the preservation of the free tips of veins R, and R,. The former of these
elements is lost in some Cylindrotominae and most Limoniini but is preserved in
virtually all Tipulinae, the Lechriini and some generalized Limoniini. The latter
element, R., is lost in many Tipulinae (as the Dolichopezaria, many Tipula) and in
the majority of the Cylindrotominae but is preserved and forms the true end of
this serial vein in many Limoniini, but not in the subtribe Limoniaria, as discussed
later. Veins R, and R, are interpreted as having fused to the margin, the resulting
vein being R,,;.

Architipulinae. The family Architipulidae, proposed by Handlirsch (1906-08,
pp. 490-491) represents at most a valid subfamily of the Tipulidae. Of the genera
placed therein by Handlirsch, only Architipula Handl. belongs here, the others
(Protipula Handl., Hotipula Handl.) being either Limoniinae or Trichoceridae.
All three genera pertain to the Upper Liassic, of an approximate age of 140,000,000
of years (Lane, 1919). The genera and species referred to the Eoptychopteridae in
the same reference are more problematical and no attempt is made herein to
correlate this material.

Tipulinae. The primitive condition of the radial field in the Tipulinae must
be assumed as being much as in the Architipulinae, that is, with cell R, deeper
than in recent genera, though approached by some species of Ctenophoraria
(Ctenophora Meig., Cnemoncosis End., and Tanyptera Latr., fumipennis O.8.,
Text-fig. 43). Other genera of this subtribe show the cell less elongate, as
Prionota v.d.W., Pselliophora O.S., Dictenidia Brullé, and some Ctenophora and
Tanyptera. The generalized condition likewise assumes that the distal sections
of veins R,, R., and R, are entire and that the true radial crossvein, 7, is present,
connecting R.,, at the fork or R, shortly beyond the origin. It should again be
emphasized that this radial crossvein is never transverse in position but always
longitudinal or oblique, forming part of the more or less zig-zag serial vein in this
subfamily. The homologies of the radial field are more obvious than in any other
group of the Tipulidae and there has been little dispute as to the correct
terminologies in this field of the wing. Comstock (1918, fig. 377; 1924, fig. 1009)
has labelled the basal section of R, as being r.

58 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

The following, chiefly Australasian genera show a primitive type of radius,
in most of them vein R, being very long and extended, greatly exceeding R.,:
Clytocosmus Skuse (skusei Alex., Text-fig. 44); Platyphasia Skuse; Ptilogyna
Westw., Phacelodocera End., Plusiomyia Skuse (felix Alex., Text-fig. 45);

R, pli R,

= R,
Rs

-R, 45

Fig. 43

Rive

Fig. 45 Fig. 46

Text-figure 43. Wing of Tanyptera fumipennis O.S.
Text-figure 44. Wing of Clytocosmus skusei Alex.
Text-figure 45. Wing of Plusiomyia felix Alex.
Text-figure 46. Wing of Leptotarsus macquarti Guer.

Ozodicera Westw., Leptotarsus Guer. (macquarti Guer., Text-fig. 46); Semnotes
Westw., Acracantha Skuse; Ischnotoma Skuse; Habromastix Skuse; Phymatopsis
Skuse; Macromastiz O.8.; Hudsonia Edw.; Zelandotipula Alex.; Longurio Lw. and
Tipulodina End.

In some species of Acracantha, Ischnotoma, Hudsonia, and the genera of the
higher Tipularia (Prionocera Lw., Holorusia Lw.; Ctenacroscelis End.;
Pectinotipula Alex.; Tipula Linn. and Nephrotoma Meig.), r is somewhat more

2nd A Ist A Cu, M

Fig. 47

Text-figure 47. Wing of Macromastix costalis Swed.

oblique in position and cell R, is smaller, vein R, being much less extended.
Holorusia, Ctenacroscelis and Zelandotipula, moreover, have vein R,; very strongly
arcuated toward vein R,,,;, strongly constricting cell R,.

In Brachypremna O.S. (candida Alex., Text-fig. 48), Tanypremna O.S.,
Megistocera Wied. (longipennis Macq., Text-fig. 52), some Macromastiz O.S.
(costalis Swed., Text-fig. 47) and others, the free tip of R, tends to become
transverse or nearly so.

BY CHARLES P. ALEXANDER. 59

Aldrovandia End. (gesneri End., Text-fig. 53) has r and the very long basal
section of R, in oblique alignment, the distal section of R. being relatively short
and paralleling R,, R,; being very extended.

In most of the genera that have been assigned to the Dolichopezaria (Dolichopeza
Curt., Text-figs. 49-51; Oropeza Ndm.; Nesopeza Alex.; Scamboneura O.8.; Megisto-
mastiz Alex.) and other genera in the Tipularia (as Microtipula Alex.; Idiotipula Alex. ;

RR,

Fig. 49 Fig. 92 ca

Fig. 53

Fig. 51 Fig. 34 Rass

Text-figure 48. Wing of Brachypremna candida Alex.
Text-figure 49. Wing of Dolichopeza malagasya Karsch.
Text-figure 50. Wing of D. cinerea Macq.

Text-figure 51. Wing of D. queenslandica Alex.
Text-figure 52. Wing of Megistocera longipennis Macq.
Text-figure 53. Wing of Aldrovandia gesneri End.
Text-figure 54. Wing of Scamboneura plumbea Alex.

Xenotipula Alex.; Tipula (Acutipula) omissinervis de Meij., and numerous species
of the genus Tipula, chiefly in the Holarctic Region) the distal section of R, is
entirely or partly atrophied. When the former is the case, as in many Dolichopeza
(queenslandica Alex., Text-fig. 51) and others, the resulting vein that connects
R, and R, is a serial vein, the cephalic portion being r, the usually much shorter
posterior portion the basal section of R, In other cases, as the subgenus
Trichodolichopeza Alex., the alignment is not so perfect. In D. malagasya Karsch
(Text-fig. 49), the basal section of R, forms an acute angle with 7 and the basal
portion of the distal section of R, is preserved as a short stub. A less accentuated
case is found in some specimens of Dolichopeza (cinerea Macq., Text-fig. 50) and
many of the species of Tipula, above mentioned.

60 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

The radial sector, short and transverse in genera such as Dolichopeza (Text-
figs. 50-51) and Nephrotoma, reaches its minimum size in Scamboneura O.S.
(plumbea Alex., Text-fig. 54) where it has apparently migrated distad beyond
the level of the other elements of the cord so that it lies at the proximal end of
the stigma and appears to be a short supernumerary crossvein; R,,, is then given
off, paralleling vein R, and forming the caudal margin of the stigma, the tips of
veins R,, R, and R,; are not especially abnormal but are greatly reduced in size
and much crowded. The basal section of R,,; is lengthened and directed strongly
basad to form the cephalic portion of the cord. There is a strong spur, sometimes
at the angle of R,,, and 7-m, in other cases on r-m itself, jutting basad into cell
R. There is an alternative interpretation to the above that is not as satisfactory
since it does not explain the small transverse element above interpreted as being
the radial sector. This interpretation would call for an elongated sector that is
almost entirely atrophied except for the small spur that juts basad into cell R,
as described. It is probable that more conclusive evidence will not be long in
coming as material in the genus Scamboneura is not especially rare in certain
Malayan islands, especially the Philippines.

In Ptilogyna, vein R,,; fuses with M,,., obliterating 7-m.

Cylindrotominae. The primitive type of the Cylindrotominae may be assumed
as having been not unlike the fossil genus Cyttaromyia Scudd. (1894, p. 190), with
the retention of the distal sections of veins R, and R, that have been lost in
Cyttaromyia and many of the living species of the group. When the fauna of
the world is considered, several species are found that have retained one or the
other of these two branches. An earlier interpretation of the venation of the
Cylindrotominae (Needham, 1908) called for a long fusion of the veins back from
the wing-margin. The true interpretation of this field was given by Osten Sacken
(1869, pp. 290-291) and later amplified by the writer (Alexander, 1914, pp. 604-605:
1919, p. 863). Brunetti (1918, pp. 280-283) has discussed the problem of the
venation in the group but has failed to appreciate the true homologies of the veins
(as is Shown by his still calling the basal section of R, in Phalacrocera replicata
the marginal crossvein, 7, and the distal section of R, the ‘costal crossvein’’).
Brunetti has summed up his beliefs in the following words: “I contend it must
be admitted that the ist vein (R,) normally and actually ends in the second
(base of R, or R,,,) because whenever the costal crossvein (distal section of R,)
is absent it most obviously does so end’. The venational symbols in parentheses
are supplied by the present writer. There can be little question but that the
Cylindrotominae have been derived from some Tipuline ancestor in middle or
late Mesozoic times.

The following modifications in the various genera may be noted: Cylindrotoma
Macq. (nigriventris Lw., Text-fig. 55) retains the primitive condition of the group
except that the distal section of R. is entirely atrophied. In a few species (as
splendens Doane), the distal section of R, is only weakly preserved or even
atrophied, though present in most individuals. Phalacrocera Schin. is of especial
importance in that the more generalized species (replicata Linn., Text-fig. 56)
have retained all three branches of this region of the wing, although R, is
represented only by a weak spur, r in this case being very long as compared with
other species of the genus. In more specialized cases (tipulina O.S., Text-fig. 57,
occidentalis Alex.) the tip of R, is entirely atrophied. In P. formosae Alex., the
tips of both R, and R, are quite atrophied. Liogma O.S. has the genotype,
nodicornis O.S., with the tips of both R, and R, atrophied. The other species agree

BY CHARLES P. ALEXANDER. 61

in this respect but some specimens of glabrata Wied. retain R, as a weak element.
In Triogma Schin., R, is strongly preserved in exculpta O.S., weakly preserved
in trisulcata Schumm., lacking in kwwanai Alex.

In Stibadocera End. (metallica Alex., Text-fig. 58), R.., diverges widely from
R,,,; and is shorter to much shorter than R,; alone. The distal section of R, is
preserved but that of R, is entirely atrophied. Stibadocerella Brun. (= Agastomyia
de Meij.) shows no trace of the distal sections of either R, or R,. Stibadocerodes
Alex. (tasmaniensis Alex., Text-fig. 59) shows no vestige of the distal section of
R,, the extreme base of the distal section of R, being more or less preserved as a

Fig, 59

Text-figure 55. Wing of Cylindrotoma nigriventris Lw.
Text-figure 56. Wing of Phalacrocera replicata Linn.
Text-figure 57. Wing of P. tipulina O.S.

Text-figure 58. Wing of Stibadocera metallica Alex.
Text-figure 59. Wing of Stibadocerodes tasmaniensis Alex.

short spur, that, when present, is provided with macrotrichiae. The fusion of
R..3 is very long, exceeding R, alone, and the space between Sc, and the basal
section of R. (the combined penultimate section of R, and 7) is correspondingly
lengthened; r-m unites with Rs before (australiensis Alex.) or at the fork.

In a few genera of Cylindrotominae, R,., has swung caudad and fused for a
greater or less extent with M,,., obliterating 7-m (some Liogma, Triogma).

It may be noted in concluding this brief account of the subfamily that the
more primitive members of the group (Cytlaromyia, Cylindrotoma) may provide
the explanation of the fate of the posterior fork of the sector in this group of
subfamilies. As has already been noted this has been called a fusion of veins
R, and R; to the margin. There is a possibility, however, that the branch inter-
preted as being M, may represent R,; of the primitive type. In Cyttaromyia this
is always connected strongly with R by a crossvein or apparent crossvein at near

62 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

midlength of the cell and in many examples of Cylindrotoma this position is still
indicated by a strong spur at the angle of M,. It is not impossible that in the
progressive outward fusion of R, and R,; in this group of subfamilies that the
latter vein in the ancestral type was deflected caudad and became fused or
intimately connected with M.

Lechriini. This new group includes the genera Lechria Skuse and
Trichoneura Lw. Lechria was placed in the tribe Hriopterini by Skuse despite
the presence of distinct tibial spurs. Later, the genus was removed to the
Hexatomini (de Meijere, Alexander). Tvrichoneura was described from the Baltic
Amber (Lower Oligocene) but has since been found living in the Himalayas.

Lechria (sublaevis Alex., Text-fig. 60) has 7 long, gently arcuated, connecting
with R, some distance from the origin of the latter. UL. singularis Skuse is figured
by its describer as having r connecting with R.,, immediately at the fork but a
photograph of the wing of the type kindly made for me by Mr. John Shewan shows

R, Sf

Fie00u Fig. 61

Text-figure 60. Wing of Lechria sublaevis Alex.
Text-figure 61. Wing of Trichoneura vulgaris Lw.

that this species, like the above, has r connecting with R, shortly beyond the
origin, forming a short basal section. The r-m crossvein is on Rs far before the
fork.

Trichoneura (vulgaris Lw., Text-fig. 61) presents a more normal Hexatomine
appearance, 7-m being connected with R,,, shortly beyond the origin.

Until the present writing both of the above genera have been referred to the
Hexatomini. The similarity of the venation to other Hexatomine genera must be
explained by convergence. An entirely comparable case is found in the genus
Trichocera Meig., which until very recently was placed with Limnophila in the
Hexatomini, the venation of the two groups being entirely comparable. It is now
known that Trichocera represents an entirely distinct family of Diptera
(Alexander, 1926).

As was indicated under the account of the lEHriopterini, the genera
Ceratolimnobia, Xipholimnobia and Xenolimnobia discussed at that time may
correctly pertain to the present group.

Limoniini. In the tribe Limoniini, the elements of the serial vein have
become straightened out in perfect alignment or nearly so. In its typical
condition, as shown by the subtribe Orimargaria, the venation of the radial field
is not difficult to homologize with the entirely similar conditions obtaining in the
Tipulinae, Cylindrotominae or the Lechriini, previously considered. In the
higher subtribes, especially in the Limoniaria, a very puzzling modification is

BY CHARLES P. ALEXANDER. 63

brought about in the various species by an outward shifting of the free tip of
R,, a condition that is discussed in detail under the account of the subtribe.

Some of the more aberrant subtribes that were formerly arranged in the
old tribe Antochini before its partition are placed in this tribe rather arbitrarily
while awaiting more data to determine their final disposition. Such genera are
Antocha, Orimargula, Orimarga, Diotrepha, Thaumastoptera, Elliptera, Helius,
Toxorhina and Dicranoptycha. The position of Toxorhina in this tribe is most
‘in doubt. Of the other generic groups placed by Osten Sacken in the Antochini
(as Limnobina anomala), Teucholabis has already been’ discussed under the
Eriopterini, and Atarba and Elephantomyia under the Hexatomini. Professor
Rogers has recently reared Atarba and some features of its life-history point
strongly to the present tribe but the affinities of the genus with Jschnothriz are
very marked and the group requires further study. The true status of many of
these aberrant subtribes of the Limoniini will probably remain long in dispute
and will be solved finally only by a knowledge of all the stages and, perhaps,
the discovery of other and annectant types.

Orimargaria. This subtribe includes Orimarga O.S. and Diotrepha O.S., the
two genera being closely allied and showing entirely comparable modifications of

Rae

x

Fig. 62 2
“a
~ R, if
=

Fig. 66

Fig. 08

Text-figure 62. Wing of Orimarga excessiva Alex.
Text-figure 63. Wing of O. punctipennis Alex.

Text-figure 64. Wing of Diotrepha fumicosta Alex.
Text-figure 65. Wing of Orimarga joana Alex.
Text-figure 66. Wing of Diotrepha atribasis Alex.
Text-figure 67. Wing of Thawmastoptera calceata Mik.
Text-figure 68. Wing of Tonnoiromyia tasnaniensis Alex.

64 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

the radial field. In many respects the group is more generalized than the
Limoniaria, despite the great modifications that have taken place in the medial
field and the recession of the m-cw crossvein. The radial sector is very long,
strongly arcuated to feebly angulated at origin, in O. punctipennis Alex. (Text-
fig. 63) nearly straight, the origin angulated and weakly spurred. The radial
field shows a remarkable plasticity of retention and loss of the distal elements.
Several species (0. excessiva Alex., Text-fig. 62; O. punctipennis Alex., Text-fig. 63;
O. andina Alex.; D. atribasis Alex., Text-fig. 66; D. fumicosta Alex., Text-fig. 64;
D. flavicosta Alex.) have retained the distal section of R, as a nearly transverse
to slightly oblique spur that is sometimes very faint and, when lost, evidently
disappears through atrophy. The distal section of R, is entirely preserved and
very long in O. excessiva (Text-fig. 62); O. pallidibasis Alex.; O. alpina Zett.;
O. dampfi Alex.; O. formosicola Alex.; D. fumicosta (Text-fig. 64); still shorter
in O. joana Alex. (Text-fig. 65); O. niveitarsis Alex.; D. mirabilis O.S., and others.
In some species this tip of R. is atrophied but leaves a spur: (0. punctipennis,
Text-fig. 63). In various species of Diotrepha, as atribasis (Text-fig. 66) the entire
distal section of R, is atrophied. The 7-m crossvein is usually not far from the
elements that constitute the cord except in the Australian O. joana (Text-fig. 65)
where it lies far distad, a condition that is perhaps explained correctly in the
original characterization of the species. The radial crossvein, 7, varies greatly
in its extent in the various species (Text-figs. 62-66).

Thaumastopteraria. The subtribe includes only the genus Thaumastoptera
Mik, with the genotype, calceata Mik (Text-fig. 67) and two extinct species from
the Lower Oligocene of Northern Europe. The group is obviously close to the
Orimargaria but the habits of the immature stages of the two groups are very
different and there are certain well-defined correlated structural characters. The
radial sector is moderately long, in alignment with R.,, and the long R,, these
three elements forming a gently sinuous curve to the wing-margin; basal section
of R,., strongly arcuated to weakly angulated; R, close to the basal section of R., r
being very short; distal section of R, entirely atrophied.

Tonnoiromyaria. The subtribe includes only the genus Tonnoiromyia Alex.
(tasmaniensis Alex., Text-fig. 68). The radial sector is long, nearly straight,
diverging gently from R,, in approximate alignment with R,,;; basal section of
R, lying distad of the level of 7-m, almost in alignment with R,, r being very
reduced and the distal section of R, entirely atrophied.

Limoniaria. The various modifications in the present subtribe, which is one
of the largest groups of crane-flies, are very complicated and are discussed in some
detail in this paper. The primitive condition of the radial field is found in a
considerable range of genera, as Libnotes Westw., Peripheroptera Schin.,
Thrypticomyia Skuse, and others. The condition in Dicranomyia (Thrypticomyia)
doddi Alex. (Text-fig. 78) may be discussed as representing this generalized
condition, in many respects not being greatly different from the Orimargaria, as
described. No member of the subtribe is known to the writer in which the distal
section of vein R, reaches the wing-margin, as in several Orimargaria, but the
extreme tip is atrophied, as shown in the various figures (Text-figs. 70-78). The
distal section of R, is preserved and lies far before the basal section of R,, 7 thus
being relatively long. The penultimate section of R,, 7 and the long distal spur of
2, are all in longitudinal alignment.

The modifications in the subtribe have been brought about in a manner that
is illustrated by actual species and may be discussed by means of the following

BY CHARLES P. ALEXANDER. 65

eight diagrams (Text-fig. 69, 1-8) arranged to show the apparent evolution of ~he
types of venation found in this subtribe.

R,
R, r R, M TR,

R, fr R, Ries RR,
6 a
AR, _R,
l ~
Rr ——— Le h
jp
Ries

Fig. 09

Text-figure 69. Diagrammatic series to show the evolution of the radial field in
the lower Limoniini. :

1. The type found in the Orimargaria: R, preserved, 7 elongate, the distal section
of R, complete, attaining the wing-margin.

2. A further development of Type 1. Note that the free tip of R, has moved distad,
materially shortening 7; distal section of R, still entire.

8. Condition as in Type 2 but the tip of R, is atrophied back from the margin.
Found in many Orimargaria, Limoniaria.

4. An accentuation of Type 3. Note that the atrophy of R, is greater and r even
more shortened. The result of this is to tend to bring the free tip of R, into approximate
transverse alignment with the basal section of R, + r. Found in numerous Limoniaria.

5. A further development of Type 4, where the atrophy of the distal section of Res
complete and the free tip of R, is in perfect transverse alignment with the combined basal
section of R, + r. This is the commonest type in the Limoniaria, being found in most
Geranomyia Hal., Rhipidia Meig., Dicranomyia Steph., some Limonia Meig., and others.

6. A type that reverts back to type 3 with a long spur of the distal section of R,
persisting ; the free tip of R, has continued its migration distad beyond the level of r and
now lies part way out on this spur. Condition found in Limonia tristigmata Alex.,
Peripheroptera schineri O.S., and several Libnotes, as greeni, Edw., notatinervis Brun.,
regina Alex., and others.

7. A further modification of type 6 where the free tip of R, has migrated to the
end of the spur of R, but still forms a rectangular bend or even with a tiny spur on R,
beyond it. In Libnotes, as wndulata Mats.

8. The culmination of the series, where the free tip of R, has migrated to the
extreme tip of the spur of R, and bends to the costa at a gently oblique angle. It should
be noted that in this case of the Limoniaria, as in all others here shown beyond type 2,
only the free tip of R, reaches the costal margin. Found in Discobola O.S., a great
section of Limonia, and other genera.

G

66 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

A few of the cases in this largest and most widespread group of crane-flies
may be further discussed:

- Libnotes Westw. In the genotype, thwaitesiana Westw., r is.very long,
meeting the basal section of R, in a long gentle curve. In several species of the
genus, a condition like type 3 of the diagrammatic series is found (as ZL.
subaequalis Alex., Text-fig. 70), being further modified in other species to type 4
(L. obliqua Alex., Text-fig. 71). Most of the species fall under type 5. Type 6
is found in L. greeni, regina and other species. Type 7 is found in L. undulata.
Thus in this single genus, we find an unbroken series illustrating five of the
diagrammatic types previously discussed.

Limonia Meig. In this genus we find a considerable range in the types, some
of which were very puzzling to the writer until the true solution of the venation
in this subtribe was finally ascertained. A great many species fall under type 5,
with the tip of R, and the basal section of R, in transverse alignment (as in
parvistigma Alex., Text-fig. 72; the Huropean annulus Meig.; bifasciata Fabr.;

Fig. 72

Re R,

Fig, 73

Text-figure 70. Wing of Libnotes subaequalis Alex.
Text-figure 71. Wing of L. obliqua Alex. ,

Text-figure 72. Wing of Limonia parvistigma Alex.
Text-figure 73. Wing of L. tristigmata Alex.

Text-figure 74. Wing of L. fumosa Alex. °

Text-figure 75. Wing of Discobola australis Skuse.

Text-figure 76. Wing of Geranomyia picta Skuse.

Text-figure 77. A. Wing section of G. tonnoiri Alex.; B. Wing
section of G. victoriae Alex.

BY CHARLES P. ALEXANDER. 67

quadrinotata Meig.; the Nearctic californica O.S.; cinctipes Say; fallax Johns.;
immatura O.S.; solitaria O.S.; triocellata O.S., and others). The other common
type in the genus is No. 8, which, at first sight, looks like a long, free distal section
of R,, with the free tip of R, atrophied (compare diagrams 1 and 8). There are
numerous representatives of this type in the Holarctic fauna (as the Huropean
flavipes Fabr.; inusta Meig.; nubeculosa Meig.; tripunctata Fabr.; trivittata
Schumm.; the Japanese amabilis Alex.; amatriz Alex.; atridorsum Alex.; basispina
Alex., esakii Alex. and many others; the Nearctic indigena O.S.; maculicosta Coaq.;
parietina O.S.; sciophila O.S.; tristigma O.S., and numerous additional species).
In two closely allied New Zealand species, LZ. sponsa Alex. falls between types
4 and 5, while L. tristigmata Alex. (Text-fig. 73) has progressed to type 6. In two
Neotropical species (L. fumosa Alex., Text-fig. 74; L. insularis Will.), R, bends
abruptly down to R,,,, a long, subperpendicular branch that is interpreted as being
R, thence extending to costa.

Discobola O.S. In the numerous Australian and New Zealand species of
this genus (australis Skuse, Text-fig. 75) type 8 obtains, the distal serial element
being very long and gently arcuated, shortest in D. tesselata O.S.

Peripheroptera Schin. The more generalized species (nitens Schin.; eudorae
Alex.) fall in type 3, with r long to very long. In other species, 7 becomes more
shortened until in some (as incommoda O.S.) type 5 is attained.

Dicranomyia Steph. The great majority of the very abundant species fall
in type 5, in a few (as punctipennis Skuse, multispina Alex.; tarsalba Alex., and
others) with the distal serial element longer than in type 5 but shorter than 8,
gently curved to the costal margin. In the subgenus Jdioglochina Alex., r tends =o
be very elongate, arcuated into the short basal section of R, The subgenus

Fig. 78

Text-figure 78. Wing of Dicranomyia (Thrypticomyia) doddi Alex.

2nd A Ist A Cu,

Thrypticomyia Skuse (doddi Alex., Text-fig. 78) has been discussed under the
introduction to the subtribe. In Huwglochina Alex. the entire radial field beyond
the cord is crowded into the wing-tip, occupying the distal fifth or less of the wing;
type 5 of the series usually obtains.

Rhipidia Meig. In the numerous species of this genus, type 5 is almost
universal.

Geranomyia Hal. The Australian species of this very large genus show a
progressive modification from a type shortly beyond 4 (with the spur of the
distal section of R, atrophied but with a distinct angulation, showing the point of
union of r and the basal section of R.; G. picta Skuse, Text-fig. 76) to other
cases that equal type 5. In G. tonnoiri Alex. (Text-fig. 77 A) R, is virtually in
alignment with the base of R, but r is still evident as a slight angulated portion
near the origin of the distal section of R,. In G. victoriae Alex. (Text-fig. 77 B),
r and the basal section of R. are in perfect alignment, and likewise about in a

68 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

transverse line with the distal end of R,. G. tridens Brun. is figured by its author
as having the distal section of R, retained, being intermediate between types 3
and 5.

Supernumerary crossveins are rather frequently found in this field of the
wing; in cell R, in the subgenus Neolimnobia Alexander (of Dicranomyia; type,
diva Schin.); in R, in Dapanoptera Westw.; in both cells R; and R, in Laosa Edw.
and Libnotes regalis Hdw.

Antocharia. This subtribe includes Antocha O.S. and Orimargula Mik. In
both, Se and R are closely approximated and the outer portion of R, lies close to
the strongly incrassated costa. Rs is straight to very gently convex, tending to be

R,.;

Rs

R,..

R,,;

Fig. 82 Ries

Rs

r-m

Fig.83 tind WRiRSD fi

Text-figure 79. Wing of Antocha saxicola O.S.
Text-figure 80. Wing of Orimargula australiensis Alex.
Text-figure 81. Wing of Hlliptera tennessa Alex.
Text-figure $2. Wing of Amphilimnobia leucopeza Alex.
Text-figure 83. Wing of Dicranoptycha germana O.S.
Text-figure 84. Wing of Heliws venustus Skuse.
Text-figure 85. Wing of Hurhamphidia viveitarsis Skuse.
Text-figure 86. Wing of Ceratocheiluws edwardsi Alex.
Text-figure 87. Wing of C. australasiae Alex.
Text-figure 88. Wing of C. flavirostris Alex.
Text-figure 89. Wing of Tozorhina magna O.S.

BY CHARLES P. ALEXANDER. ~ 69

in alignment with R,,;; basal section of R. preserved, the distal section very short.
In Orimargula (australiensis Alex., Text-fig. 80), veins Sec, and R, are more
nearly approximated at the wing-margin than in Antocha (sazicola O.S., Text-
fig. 79).

Ellipteraria. The subtribe includes only the genus Hlliptera Schiner. Rs is
elongate, lying very close to and paralleling R,, in alignment with R.,,; basal
section of R. preserved in some species (tennessa Alex., Text-fig. 81): jacoti Alex.;
zipanguensis Alex.), placed at or shortly beyond the fork of Rs, the section lost by
atrophy in other species (omissa Egg.; clausa O.S.; astigmatica Alex., and others) ;
R, elongate, in alignment with R.,,; basal deflection of R,,; very strongly arcuated
to angulated and spurred (illini Alex.) or nearly straight and transverse (jacoti).

Amphilimnobiaria. The single genus and species, Amphilimnobia leucopeza
Alex. (Text-fig. 82) is included. Rs is long, angulated and short-spurred at origin:
basal section of R. preserved, lying just proximad of the level of r-m and opposite
the tip of Sc.

Dicranoptycharia. The single genus Dicranoptycha O.S. (germana O.S., Text-
fig. 83) is included. Rs is variable in length, the veins at its fork symmetrical;
R,,, very elongate, the basal section of R, lying far distad, the distal section of R.
being very short.

Heliaria. Rs is short and often straight. In Hurhamphidia Alex. (niveitarsis
Skuse, Text-fig. 85), r-m lies far before the fork of Rs. Rhampholimnobia Alex.
is similar but r-m lies closer to the fork. In Helius (venustus Skuse, Text-fig. 84),
r-m is usually on R,,; shortly beyond its origin. In many species of the genus, 7-m
is obliterated by the fusion of vein R,,; on M,,... In H. wniformis Alex., r-m is
opposite the outer end of cell ist M,. In all genera and subgenera of this
subtribe, the basal section of R, is entirely atrophied.

Toxorhinaria. This subtribe includes the two genera Ceratocheilus Wescheé
and Toxzorhina Lw. The strict position of the subtribe is still much in doubt and
it is placed in the Limoniini with certain misgivings. In the more generalized
species of Ceratocheilus (edwardsi Alex., Text-fig. 86; qaustralasiae Alex., 'Text-
fig. 87), the basal section of R. appears to have been lost as in the preceding sub-
tribe, the long distal section of R, reaching the margin in a gently sinuous curve.
In still other species of the genus (flavirostris Alex., Text-fig. 88) a progressive
shortening of the branch R, takes place until it has shortened into a small and
nearly transverse element. In Toxorhina (magna O.S., Text-fig. 89) this vein R,
is entirely lost but whether by final fusion of the element with R. (as previously
discussed for the Hexatomine genus Psaronius and the Eriopterine genus
Gymnastes) or by the atrophy of R, when it has become short and weak cannot
be stated without more evidence.

PHYLOGENETIC CONSIDERATIONS.

The two principal divisions of the Tipulidae have been distinct since at least
the middle of the Mesozoic (Upper Liassic), at which time the Architipuline and
Tanyderid types co-existed. From the Architipulinae, the Tipulinae (Text-fig. 90)
may be derived in almost a direct line. The Cylindrotominae are quite as
obviously an offshoot of the Tipulinae. The two Limoniine tribes, Limoniini and
Lechriini, that appear to have been evolved from this same general strain have
been isolated from the above and from one another for a longer time and it does
not seem possible to derive them directly from the Tipuline stem.

70 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

The Pediciini have been derived from some pre-existing type that was not
conspicuously unlike the recent Tanyderidae. From the Pediciini, through the
subtribes Adelphomyaria and Ularia, the tribe Hexatomini is derivable, leading,
in turn, through the Hexatomoid Eriopterini to the higher members of the latter
tribe. The exact point of origin of the Styringomyiini must be held in question.

STYRINGOMYIINI

NI ERTOPTER INT

TIPULINAE

CYLINDHOTOMINAE
HEXATOMINI

ULARIA RICHOCERIDAE
PEDICIINI

mm

Lv

IPTERON

Pa he

Fig. 90

Text-figure 90. Phylogenetic Tree.

The Ptychopteridae (Ptychoptera lenis O.S., Text-fig. 91) have apparently been
derived from the Psychodoid stem, but, if so, they have independently had vein R,
unite with R, to form a short fusion back from the margin, much asin the
Pediciini. The Trichoceridae (Trichocera annulata Meig., Text-fig. 92), in similar
manner, must have been derived from some primitive ancestor in this general
region yet it too has undergone the cephalization of vein R. and the capture of
vein R, by R,.,, quite as in the higher Tipulidae. It is highly probable that these

Fig. 9]

Text-figure 91.
Text-figure 92.

Sc, Sc,

Fig. 92

Wing of Ptychoptera lenis O.S. (Ptychopteridae).
Wing of Trichocera annulata Meig. (Trichoceridae).

BY CHARLES P. ALEXANDER. — (al

two families have led more or less directly to the more specialized groups of
Nematocera and thence to the Brachycera, and the correct interpretation of the
basal section of R. becomes of critical importance in any attempt to trace the
phylogeny and correct homologies of the higher Diptera.

Of the other families of Nematocera, the Culicidae, Dixidae, Chironomidae,
Orphnephilidae, Anisopodidae, Blepharoceridae, Simuliidae and Bibionidae all
agree with the interpretation of the wing, as given, except that the basal section
of R. in all cases has atrophied and in all except the most generalized forms R,
and R, have fused to the wing-margin. The other Nematocerous groups, with a
more highly specialized venation, as the Deuterophlebiidae, Scatopsidae, Sciaridae
and Cecidomyiidae offer little direct evidence from the few veins that are still
retained in the radial field. From other structures, however, the affinities of the
groups with the others above listed render it necessary to interpret the venation as
is done in the latter case.

The lower groups of the Orthorrhaphous Brachycera in similar manner agree
entirely with the above interpretation, veins R, and R; remaining distinct as in
the Ptychopteridae. From these phylogenetically lower families of Brachycera,
the higher groups of the Orthorrhapha and the Cyclorrhapha have been derived.

Summary.
The main points of the preceding discussion may be summarized as follows:

1. The so-called radial crossvein (r) in all Diptera except a limited group of
the Tipulidae (Tipulinae, Cylindrotominae) is, in reality, the basal section of vein
R. which has swung cephalad across cell R, and fused back from the wing-margin
with vein R,, forming a short or longer fusion, R,,».

2. There are two distinct lines of specialization in the Tipulidae that have
been distinct since at least mid-Mesozoic times. In the first, represented by the
Limoniine tribes Pediciini, Hexatomini and Eriopterini, the anterior branch of
the upper fork of the radial sector (R.) of the primitive Tanyderoid ancestor has
swung cephalad and fused with R,, as discussed under 1; by a slight shifting of
the veins at the end of the sector, the anterior branch of the lower fork of the
sector (R,) becomes more intimately attached to the stem of the upper fork (R.,,)
forming a short to longer fusion of veins R,,,,,; in higher groups, veins R, and R,
may become fused to the wing margin, obliterating cell R;. The basal section
of R, is present in most cases but in many scattered groups has atrophied, leaving
no indication of its previous position. The posterior branch of the sector is thus
R; alone. In the second group, represented by the Architipulinae, Tipulinae,
Cylindrotominae, and the Limoniine tribes Lechriini and Limoniini, the true radial
crossvein (7) is present but always in a longitudinal or oblique position, forming
part of a more or less perfect serial radial vein. In this group, the distal sections
of R, and R, are lost by atrophy. The posterior branch of the sector is inter-
preted as being a fusion to the margin of veins R, and R;.

3. In some few Pediciini, of the group Dicranotae, there is a supernumerary
crossvein in cell R, that simulates a true primary crossvein in this cell.

References.
ALEXANDER, C. P., 1914.—New or little-known craneflies from the United States and
Canada. Tipulidae, Diptera. Proc. Acad. Nat. Sci. Philadelphia, 1914, pp. 579-695.

, 1918.—A new interpretation of the wing-venation of the Pediciine crane-flies
(Tipulidae, Diptera). Ent. News, xxix, pp. 201-205.

72 INTERPRETATION OF THE RADIAL FIELD OF WING IN DIPTERA.

, 1919.—The crane-flies of New York. Part i. Distribution and Taxonomy of
the Adult Flies. Cornell Univ., Agr. Expt. Sta., Mem. xxv, pp. 860-869.

, 1920.—New or little-known crane-flies from Formosa (Tipulidae, Diptera).
Ann. Ent. Soc. America, xiii, pp. 249-270.

, 1926.—The Trichoceridae of Australia (Diptera). Proc. LINN. Soc. N.S.W.,
li, pp. 299-304.

BRUNETTI, H., 1918.—Revision of the Oriental Tipulidae with descriptions of new species,
part ii. Rec. Indian Mus., xv, pp. 255-340.

Comstock, J. H., 1918.—The wings of insects, pp. 1-430.

, 1924.—An introduction to entomology, pp. 1-1044.

Comstock, J. H., and NEEDHAM, J. G., 1898-99.—The wings of insects. Amer. Nat., xxxii,
43, 81, 2381, 237, 240, 248, 249, 253, 256, 335, 413, 420, 423, 561, 769, 774, 903 (1898) ;
XXXIll, 118, 573, 845, 851, 8538, 858 (1899).

CRAMPTON, G. C., 1924.—Remarks on the phylogeny and interrelationships of Nemato-
cerous Diptera. Psyche, xxxi, pp. 238-242.

, 1925a—A phylogenetic study of the thoracic sclerites of the non-Tipuloid
Nematocerous Diptera. Ann. Ent. Soc. America, xviii, pp. 49-74. A

, 1925b.—Evidences of relationship indicated by the thoracic sclerites of certain
Hriopterine Tipuloid Diptera. Jnsec. Inscit. Menst., xiii, pp. 197-213.

, 1926a.—A phylogenetic study of the thoracic sclerites of the Psychodoid Diptera,
with remarks on the interrelationships of the Nematocera. Ent. News, xxxvii, pp.
33-39, 65-70.

———, 1926b.—The external anatomy of the primitive Tanyderid Dipteran Wacrochile
spectrum Loew, preserved in Baltic Amber. Bull. Brooklyn Ent. Soc., xxi, pp. 1-14.

EpWarps, F. W., 1921.—British Limnobiidae. Some records and ccrrections. Trans. Hint.
Soc. London, 1921; pp. 196-230.

, 1926.—The phylogeny of Nematocerous Diptera: a critical review of some

recent suggestions. Third Internat. Ent. Congress, 2, pp. 111-130.

HANDLIRSCH, A., 1906-08.—Die Fossilen Insekten, pp. 1430.

JOHNSON, C. W., 1901.—Variation in the venation of Amalopis inconstans Osten Sacken.
Ent. News, xii, pp. 305-307.

LAMEERE, A., 1906.—Notes pour la classification des Dipteres. Mem. Soc. Entom.
Belgique, xii, pp. 105-140.

LANE, A. C., 1919.—Geological Column: France, Britain, Germany, United States. Dewey
Lefax System, Dewey 550, Lefax 51, pp. 9-357-359.

Lorw, H., 1862.—Monographs of the Diptera of North America, Part i, x-xxiv.

MacGi.uivray, A. D., 1923.—External Insect-Anatomy, pp. 1-388.

NEEDHAM, J. G., 1908.—Venation of the wings of Tipulidae. 237d Rept. State Ent. New
York, 1907, pp. 217-238. ;

OsTEN SACKEN, C. R., 1869.—Monographs of the Diptera. of North America. Part iv,
pp. 1-345.

ScHINER, J. R., 1862.—Fauna Austriaca. Die Fliegen (Diptera), 1, pp. 1-672.

ScupprEr, S. H., 1894.—Tertiary Tipulidae, with special reference to those of Florissant,
Colorado. Proc. Amer. Philosoph. Soc., xxxii, pp. 163-245.

TILLYARD, R. J., 1919.—The Panorpoid Complex. Part 38. The wing-venation. Proc.
LINN. Soc. N.S.W., xliv, pp. 533-718.

WILLISTON, S. W., 1908.—Manual of North American Diptera, Ed. 3, pp. 1-405.

THE ANATOMY OF CHEILANTHES VELLEA.

By May M. ‘Witiiams, M.Sc.
Linnean Macleay Fellow in Botany.

(Twenty Text-figures. )
[Read 30th March, 1927.]

Introduction.

In a previous paper (Bot. Gaz. Ixxviii, No. 4, 1924), the writer described the
anatomy of Cheilanthes tenuifolia. The interest of this type lies in the fact that
the rhizome possesses a true dictyostelic structure, while the petiolar stele consists
of a single strand. The xylem elements of the latter contain three protoxylem
groups, the median one of which varies in position until it finally becomes exarch
in position and then completely detached from the metaxylem elements. Since
there is so much variation of stem and petiolar anatomy in the various species of
the genus Cheilanthes, the investigation of Ch. vellea (one of the other two species
common in New South Wales) was undertaken for the purpose of comparison with
Ch. tenuifolia and other foreign types.

Marsh (1914) described four American species namely Ch. persica, Ch. fendleri,
Ch. gracillima and Ch. lanuginosa. Gwynne-Vaughan (1903) described the stelar
anatomy of Ch. lendigera and Ch. microphylla. All these types show considerable
divergence of stelar structure, but, at the same time, form an excellent series
for studying the gradual transition from solenostely to dictyostely. Prantl (1882)
described the development of the young sporangia of Cheilanthes and considered
that as the young sporangia develop quicker than the margin, the appearance is
almost as though they sprang directly from the marginal cells.

The various species of the genus Cheilanthes are mostly small ferns with
short horizontal rhizomes. This is the case in Ch. vellea: the external morphology
shows various adaptations to the xerophytic habitats. The fronds are stiff and
pinnately compound: the lower side is often brownish and always covered with a
dense growth of hairs. The pinnae are usually opposite, in Ch. tenwifolia they
are alternate. The sori are continuous and protected by the inrolled margin of
the leaf. Ch. vellea is common in the dry parts of the interior of New South
Wales.

The material for the present investigation was collected at Wiseman’s Ferry
(1923) and at Pennant Hills (1926). The material was fixed in a 1% chromo-
acetic solution, washed, dehydrated and passed in the usual manner into paraffin.
Sections were then cut 5 w to 12 uw in thickness: Flemming’s Triple stain, also
Safranin and Haematoxylin were the staining processes most extensively employed.

The anatomy of Ch. vellea very closely resembles that of Ch. tenuwifolia with

various differences principally in the stelar structure of the petiole.
H

74 THE ANATOMY OF CHEILANTHES VELLEA.

Investigation.
A. Rhizome.
The rhizome is covered by a fairly extensive development of scales, which are

very dense around the apex. As indicated in Text-fig. 1, the apex is broad and
flat, while the apical cell arises as a small conical emergence. The apical cell
is tetrahedral in shape and segmentation is identical with that described in
Ch. tenuifolia (1924). About 1mm. behind the apex, the whole of the ground
tissue becomes sclerized.

In Ch. vellea the structure of the stem stele is of that type known as the
dictyostele, due to the overlapping of the leaf gaps (Text-fig. 2). Although the

Text-fig. 1. Longitudinal section of the apex of the rhizome. The apical cell
(X) is tetrahedral in shape and forms the typical segments. x 240.

Text-fig. 2. Transverse section of the rhizome indicating its dictyostelic
structure. Three meristeles (M) are shown and three overlapping leaf
gaps (L.G.). x 60.

BY MAY M. WILLIAMS. 75

structure is dictyostelic, it is only a simple type of such a form of anatomy as
there were never more than three or four leaf gaps present in any section
examined. Ch. tenuifolia is a more advanced type in this respect as the vascular
cylinder is more frequently broken by leaf gaps than in Ch. vellea. The stelar
anatomy of these two types at once places them near Ch. persica (Marsh, 1914)
which is also a distinctly dictyostelic form. The other types which have so far
been investigated show considerable variation of stelar structure.

Ch. fendleri (Marsh, 1914) is a true solenostele.

In Ch. gracillima (Marsh, 1914), the stele behaves as a true solenostele
except at certain points where apparently two leaves are given off together.

In Ch. lendigera and Ch. microphylla (Gwynne-Vaughan, 1903), each leaf
gap in the stele is closed before the next above is formed, so far as the endodermis
and pericycle are concerned. The leaf-gap in the xylem, however, remains open
until it overlaps with the gap formed in the xylem by the departure of the next
leaf-trace above.

In Ch. lanuginosa (Marsh, 1914), the leaf traces are given off singly, in pairs
or even in threes, so that at intervals the vascular cylinder may be a complete
ring, though more frequently it is broken giving a dictyostelic structure (cf.
Marsh, Text-fig. 14).

This series taken in conjunction with Ch. vellea, Ch. persica and Ch. tenwifolia
indicate well a gradual transition from solenostely to dictyostely.

In Ch. vellea, as in all the other types, the leaf trace comes off trom the
centre of the meristele; its separation from the stem stele is not effected
synchronously on both sides. The root trace passes off in the usual manner.

As indicated in Text-fig. 3, the meristele is composed of a central core of
xylem intermingled with which is a little xylem-parenchyma. There is no trace
of protoxylem present in the rhizome, even at the apex, all the tracheids of the
xylem having sclariform thickening on their walls. The central core of xylem
is not as wide as that present in Ch. tenuifolia, nor is there so much xylem
parenchyma present. This is precisely the condition which follows on Marsh’s
generalization, since Ch. tenuifolia has a more complicated dictyostele than that
present in Ch. vellea. Marsh (1914) found that there is a close relation between
the stelar structure and the amount and arrangement of the wood within the
stele. In solenostelic types there is a narrow continuous band of tracheids, not
broken up by parenchyma. In dictyostelic types there is a broad band of wood
among which a good deal of parenchyma occurs.

The xylem is surrounded by a layer of ordinary parenchyma (Vasal parenchym
of Strasburger), followed by a layer of phloem cells. The phloem never consists
of more than a single layer of sieve tubes, all of which are of uniform size so
that there is no differentiation into protophloem and metaphloem (Text-fig. 4).
The phloem does not continue around the leaf gap so that there is no continuity
of internal and external phloem; surrounding the phloem are two or three layers
of fairly large parenchymatous cells, which as in Ch. tenuifolia might be inter-
preted as representing a multilayered pericycle; surrounding this is the endo-
dermis. The cells of the pericycle are of interest owing to the fact that they
contain a great deal of granular matter which is always arranged at one side
of the cell, and in the same direction in each cell giving the whole structure a
peculiar, but distinctive, appearance. The granules lie on that side of the cell
which is the lower side in the position in which the rhizome is lying. The granules
are not starch granules but are evidently food granules of various kinds. This

76 THE ANATOMY OF CHEILANTHES VELLEA.

(Ole

65 ey

ee

o

Text-fig. 3. Transverse section of a meristele showing the central core of xylem (Xy.)
containing xylem parenchyma (Xy.P.), surrounded by “vasal parenchym” (V.P.), a
single layer of phloem cells (Ph.), a multilayered pericycle (P.) and the endodermis
Cen, ax) wb0:

Text-fig. 4. A portion of the above indicating the nature of the xylem (Xy.) and the
internal (I. Ph.) and external (Ex. Ph.) phloem. x 240.

Text-fig. 5. <A few of the cells of the pericycle indicating the position of the granules.
x 240.

Text-fig. 6. Longitudinal section of the apex of ihe leaf, indicating the two-sided wedge
shaped apical cell and its segmentations. x 400.

Text-fig. 7. Transverse section of the vascular strand to the petiole. The three
protoxylem groups (Pr. Xy.) are shown. x 150.

BY MAY M. WILLIAMS. : 77

condition is not due to fixation as it is to be seen in the living material. The
structure is very like that which has been described and figured by Schaefer in
various root tips. The condition is evidently a purely physiological one aud
related to gravity. (Text-fig. 5.)

The rhizome structure at once places Ch. vellea as an advanced type of the
genus.

B. Leaves.

As indicated in Text-fig. 6, the growth of the leaf is due to a two-sided, wedge
shaped apical cell forming two rows of segments, such as is typical for
Leptosporangiate Ferns. The outer divisions of the segments become the marginal
cells of the pinnae, the other divisions produce the petiole. The young leaf
invariably shows circinate vernation.

Petiole.

The petiolar stele forms a single strand surrounded by a sclerenchymatous
cortex. Text-fig. 7 indicates the vascular strand, the main mass of which is
composed of large metaxylem elements in conjunction with which appear three
protoxylem groups. The ends of the xylem mass are slightly curved inwards. This
is followed by a single layer of parenchymatous cells surrounded by a layer of
phloem cells. The phloem is continuous around the strand. This is followed by
a few more layers of parenchymatous cells—the pericycle—the whole being
enclosed by a typical endodermis. This has usually broken down in the older parts
of the petiole. The parenchymatous cells are filled with dense granular contents
and contain large nuclei.

The principal interest of the petiolar stele of the genus Cheilanthes lies in the
various positions occupied by the third median protoxylem group in the different
species and at different levels of the petiole of the same species. The vascular
strands of the rhizome show no visible protoxylem elements, but as soon as the
leaf trace leaves the stem stele, protoxylem groups make their appearance in the
trace which had hitherto been without them. As already mentioned, at the very
base the condition is that of a simple adaxially curved anarch leaf trace.
Text-fig. 8 indicates a section taken from the base of the young petiole. At this
level there are two lateral, endarch protoxylem groups, while the third median
group is distinctly exarch in position and attached to the as yet unlignified
metaxylem elements.

Text-figs. 7 and 9 indicate sections made at higher levels of the petiole. Here
the median protoxylem group has become completely detached from the metaxylem
and is separated from it by parenchyma. The number of tracheids composing this
group is also very much less usually consisting of two or three tracheids often
widely separated. At higher levels this group dies out altogether.

The condition here described can easily be brought into line with that which
occurs in the species described by Marsh (1914) and that of Ch. tenuwifolia
(Williams, 1924).

Marsh (1914, p. 678) maintains that in all the species which he investigated
there was at the base a simple adaxially curved anarch leaf trace, but as the
petiolar strand is followed upwards changes occur in the petiolar strand which vary
considerably. These changes may be briefly outlined as follows: In Ch. fendleri
the anarch strand soon develops three protoxylems all of which are endarch.
The metaxylem immediately exterior to the median protoxylem group dies out;

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THE ANATOMY OF CHEILANTHES VELLIA.

78

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Text-fig. 8. Transverse section of the young petiole near the base.

positions of the three protoxylem groups are clear. x 150.
Text-fig. 9. Transverse section of the upper portion of the petiolar stele at a higher

level. The median protoxylem group is here detached and broken up. x 240.

BY MAY M. WILLIAMS. 79

the two portions of the stele soon rejoin, but the junction is effected in such a
way that the protoxylem is now exarch in position. The protoxylem next becomes
completely detached and runs through the major portion of the petiole as a single
separate group. Later it becomes broken into three or four smaller groups.

In Ch. lanuginosa the three protoxylem groups appear at the base, but at this
point the centre of the trace is so narrow that the median protoxylem occupies
the whole of the thickness of the stele. The two metaxylem masses rejoin, but
the median protoxylem does not persist. It is found for a short distance as a
line of thickening among the parenchyma but at higher levels no trace of it can
be recognized. ‘

In Ch. gracillima the two lateral protoxylems occupying an endarch position
are the only ones which can ever be definitely recognized at any level of the
petiole.

In Ch. persica the petiolar stele has the shape of an obtuse angled isosceles
triangle, and: has the three protoxylem groups at each angle.

In Ch. tenuifolia (Williams, 1924, p. 384-385) the petiolar stele is described
as consisting of, at the base, a simple adaxially curved anarch structure. When
the protoxylem groups appear, the two lateral components take up the typical
endarch position, while the metaxylem immediately exterior to the median
protoxylem has died out, leaving the protoxylem as the sole junction between
these two masses. These, however, soon rejoin, and median protoxylem becomes
exarch. At a higher level it was found to be detached from the metaxylem
elements.

It is thus seen that Ch. tenuifolia might be interpreted as a form intermediate
between Ch. fendleri and Ch. lanuginosa with regard to the structure of the
petiolar stele, since various stages at the base of the petiole in Ch. fendleri are
not present in Ch. tenuifolia, while various phases at the apex in Ch. lanuginosa
are not represented in Ch. tenuifolia. In the same manner Ch. vellea might be
described as intermediate between Ch. lanuginosa.and Ch. gracillima for the type
of petiolar structure.

Marsh (1914, p. 680) maintains that these facts admit of but one explanation,
that is, that the base of the leaf preserves primitive characters. He considers
that it is difficult to imagine what could actually be the cause of the exarchy and
detachment of the median protoxylem: it is simple from a study of leaf base
transition to derive Ch. fendleri from the normal type and the Cheilanthes petioles
from Ch. fendleri. He also states (p. 681) that in deriving Ch. lanuginosa and
Ch. gracillima from Ch. fendleri that the earlier stages may be dropped out at
the base of the petiole. This dropping out of the earlier stages is also to be
observed in Ch. vellea and Ch. tenwifolia.

Sinnot (1911) maintains that in the triarch petiolar stele, it is always the
median protoxylem group which more readily becomes broken up into two
groups. He has also spoken of the “striking constancy” of endarchy in the
filicinean petiolar stele. The investigations recorded on the various species of
Cheilanthes illustrate very well the plasticity of the median group with its
gradual movement from an endarch to an exarch position and finally its complete
separation and disintegration. While these types mainly agree with Sinnot’s
generalizations of the filicinean leaf trace, they certainly do seem to have slipped
ahead in the general evolutionary scheme.

It is quite generally recognized that the stem and petiolar structure are
simply and fundamentally the result of an adaptation to the demands made upon

80 THE ANATOMY OF CHEILANTHES VELLEA.

Cr

SS
KS4O'S
[Be

Text-fig. 10. Transverse section of the pinnule indicating the nature of the upper and
lower epidermis, the palisade and lacunar tissue. x 150.

Text-fig. 11. A surface section of the lower surface of the pinnule, indicating the nature
of the stomata. x 150.

Text-fig. 12. Transverse section of the vascular bundle of the pinnule. x 240.

Text-fig. 13. Transverse section of the apex of the root showing the apical cell with its
typical segmentation. x 240.

Text-fig. 14. Transverse section of the root showing the sclerized cortex and the central
vascular strand. x 60.

BY MAY M. WILLIAMS. 81

these parts by the foliage. Taking this fact into consideration, it might be well
to look to the habitat of these Ferns for explanations of any peculiar types of
structure present in the stem or petiole. It will be remembered that these ferns
are typically xerophytic. Other small ferns found growing under similar xerophytic
conditions are Lindsaya linearis and L. microphylla. The writer has investigated
the anatomy of these types (1925) and observed in the petiolar steles the same
exarch positions of the median protoxylem groups. In these types, however, there
was not to be observed that transition from an endarch to an exarch position.

The tentative explanation offered for this variation in the position of the
median group is that it is a response to the unique habitat of these ferns; that is,
it is an adaptation to dry conditions.

Pinnule.

The pinnules are lobed; the veins show dichotomous venation, and on the
ends of these forked veins the sori are borne. The lower surface of the leaf is
protected by a dense mass of hairs.

Text-fig. 10 indicates the minute anatomy of the pinnule. This is very similar
to that of Ch. tenuifolia. The cells of the upper epidermis are large, are filled
with dense granular contents, and are protected by the cuticle. The lower
epidermis consists of relatively thin walled cells, and from these grow out the
dense mass of hairs which cover the lower surface. The stomata are confined
to the lower epidermis and open into large air cavities; they are protected by
the inrolled margins of the pinnule plus a covering of hairs. (Text-fig. 11.)

The lacunar tissue is reduced, while the palisade tissue is well developed,
often being three cell layers deep. The cells are oblong in shape, and the
chloroplasts are arranged, for the most part, along the margins of their walls
(Text-fig. 10). The vascular bundles of the pinnae are usually collateral. Each is
surrounded by a well defined endodermis, the cells being large, colourless and thin
walled. This encloses a layer of parenchymatous cells—the pericycle-——-which
surrounds a central mass of xylem tracheids and a few sieve tubes representing
the phloem (Text-fig. 12).

A vascular commissure carries the vascular supplies to the developing
sporangia.

From these remarks, it will be seen that the leaf, both in external morphology
and minute anatomy, shows well marked adaptations to its xerophilous habitat.

C. Root.

The roots present no features of particular importance in the present
discussion. They arise in connection with the bases of.the leaves. The vrimary
roots branch profusely; the secondary roots are produced in two lines corres-
ponding to the protoxylem groups in the vascular bundle of the primary root.

The apex is characterized by the presence of a three sided, pyramidal apical
cell, from which segments are cut off in the usual manner (Text-fig. 13).

Text-fig. 14 indicates the minute anatomy of the older root. Here the cortex
has become sclerized and surrounds a diarch vascular strand. ‘The latter is
composed of a central core of xylem, the metaxylem elements being large and
few with the protoxylem groups arranged at the ends. The phloem consists of a
single row of sieve tubes which do not continue around the protoxylem groups.
The xylem and phloem are separated by one or two layers of parenchyma.

Surrounding the phloem and separating it from the endodermis, are three or four
I

82 THE ANATOMY OF CHEILANTHES VELLEA.

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MY GAM CO:
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Text-fig. 15. Transverse section of the vascular bundle of the root, indicating its diarch
nature. x 240.

Text-fig. 16. This indicates the nature of the soral region of the pinnule. x 150.

Text-fig. 17. This indicates the nature of the first segmentation occurring in the
sporangium. x 240.

Text-fig. 18. This indicates that stage in the development of the sporangium when the
wall and stalk cells are formed and the central cell which will form the archeosporium
proper. x 240.

Text-fig. 19. The typical quadrant stage of development of the sporangium. x 240.:

Text-fig. 20. This indicates the nature of the mature spore. x 240.

BY MAY M. WILLIAMS. 83

layers of parenchymatous cells which might be interpreted as representing the
pericycle. The parenchymatous cells have dense granular contents and large
nuclei (Text-fig. 15).

D. Sorus.

The sori are superficial and continuous on the back of unthickened nerve
endings. They are protected by a false indusium which is composed of the inrolled
margin of the pinnule. The soral region is represented in Text-fig. 16. It is broad
and flat and shows the typical mixed character with regard to the arrangement
of the sporangia.

The development of the sporangia is identical with that described in Gh.
tenuifolia (Williams, 1924), hence for that typical for Leptosporangiate Ferns.
The sporangia arise from an epidermal cell which protrudes above the surface and
becomes divided by a transverse wall into a base or stalk cell and an upper cell
which forms the sporangium proper. Since development is of the general nature
typical for Leptosporangiate Ferns, there is no need to describe it here. Various
stages in the development are represented in Text-figs. 17, 18, 19.

The annulus is vertical and interrupted by the stalk. The number of spores
produced in each sporangium does not exceed 16-20. The nature of the mature
spore is shown in Text-fig. 20.

Conclusions.

The convergence of evidence seems to indicate that the genus Cheilanthes
represents a group of ferns which in the earlier types show anatomically primitive
characters, but possessing soral characters which are advanced. In some species,
the stem stele becomes more complicated by the insertion of extra leaves and the
overlapping of the gaps formed by their departure. These primitive anatomical
characters present in the stem might perhaps be interpreted in terms of the
habitat of these ferns, combined with the reduction in the foliage.

In all parts of these ferns there are striking adaptations to xerophily. Marsh
(1914, p. 677) regards the reduction in the number of sieve tubes constituting the
phloem in all parts of the plants as such an adaptation. He states that “the
phloem of these forms is remarkable for the absence of differentiation into proto-
phloem and metaphloem. . . The whole of the phloem of these forms may fairly
be said to correspond with the protophloem of mesophytic species, the slow growth,
necessitated by the xerophilous habitat rendering unnecessary those facilities for
active translocation which would be given by large metaphloem elements.” The
writer has also suggested that the structure of the petiole with the peculiar
movement of the median protoxylem group is of the nature of a response to the
unique environment of these ferns.

Summary.

1. The apex of the stem is occupied by a tetrahedral apical cell cutting off
segments parallel to the cutting faces.

2. The stem stele is of that type of anatomy known as the dictyostele; this
at once places Ch. vellea as one of the more advanced species belonging to the
genus.

3. The leaf is formed by segmentation from a two-sided, wedge shaped apical
cell. —

4. The petiolar stele as it leaves the stem stele is a simple adaxially curved
anarch leaf trace.

84 THE ANATOMY OF CHEILANTHES VELLEA.

5. When the protoxylem groups appear, the two lateral components are
endarch in position while the median group is exarch.

6. At higher levels in the petiole the median protoxylem group becomes
detached from the metaxylem elements; it becomes broken into three or four
smaller groups and finally disappears.

7. The pinnae have a reduced lacunar system and a well developed palisade.

8. The stomata are confined to the lower surface of the pinnae and are
protected by the inrolled margins of the pinnae together with a covering of hairs.

9. The vascular bundles of the pinnae are collateral.

10. The apex of the root is occupied by a three sided apical cell with typical
segmentations.

11. The root is diarch.

12. The sorus is superficial in origin and continuous. It is protected by the
inrolled margins of the leaf alone.

13. The development of the sporangia conforms with that typical for
Leptosporangiate Ferns. ;

14. Cheilanthes vellea shows, both in external morphology and internal
anatomy, well marked adaptations to its xerophilous habitat.

In conclusion, the writer wishes to express her thanks to Professor Lawson,
in whose laboratory the investigation was carried out, for advice and guidance.

Literature Cited.
Bower, F. O., 1918.—Studies in the Phylogeny of the Filicales. VII. The Pteroideae.
Ann. Bot., XXXII, 1-68.
, 1923.—The Ferns. Camb. Bot. Handbooks.
GWYNNE-VAUGHAN, D. T., 1901.—Observations on the Anatomy of Solenostelic Ferns. I.
Loxsoma. Ann. Bot. XV. 71-92.
, 1903.—ITd. II. Ann. Bot. XVII. 689-742.
MarsH, A. S., 1914.—The Anatomy of some Xerophilous species of Cheilanthes and
Pellaea. Ann. Bot. XXVIII... 671-6838.
SINNOT, E. W., 1911.—The Evolution of the filicinean Leaf Trace. Ann. Bot., XXV.
167-190. : j
WiuuiAMs, M. M., 1924.—Anatomy of Cheilanthes tenuifolia. Bot. Gaz., LXXVIII. ~
378-396.
, 1925.—The Anatomy of Lindsaya linearis and Lindsaya microphylla. Proc.
LINN. Soc. N.S.W., L. 391-404.

THE GEOLOGY OF THE COUNTRY BETWEEN LAMB’S VALLEY AND THE
PATERSON RIVER.

By G. D. Osporne, B.Sc., Lecturer in Geology, University of Sydney,
formerly Linnean Macleay Fellow of the Society in Geology.

(Plate i and two Text-figures. )
[Read 30th March, 1927.]

Introductory and Explanatory.

The geological investigation of the country described in this paper began in
1921 when Professor Browne and I made a brief examination of the sections east
and north of the Gosforth and Hillsborough district. Following this preliminary
work a number of intermittent trips were made to the area, Paterson, Vacy and
Rosebrook being used as bases at various times, while on two occasions a stay was
made at the house on Portion 42, Parish of Wolfingham, owned by Mr. W. Sherwood
of Branxton.

The main portion of the area examined consists of a somewhat dissected
plateau, and on account of the almost entire absence of settlement and of good roads
into the area, access is somewhat difficult, and much time is consumed in
travelling to the region of investigation. For these reasons I have not been able
to make the geological examination as detailed as I desired, and it is probable
that much more information concerning the detailed geological features can be
gained by paying closer attention to the region.

Sufficient work, however, was done to arrive at an understanding of the
general features of the structure, stratigraphy, physiography and petrography, and
the field-survey was such as to permit the drawing of the map given on Plate i.

In connection with the carrying out of the work I wish to record my indebted-
ness to Mrs. Hudson and family of “Hillsborough,” Mr. and Mrs. Dick Cant, Mr.
and Mrs. A. McDonald of Gosforth, and Mr. Peter Campbell and family of Rosebrook
for hospitality while visiting the area. I am also grateful to Mr. W. Sherwood
for permission to stay on his property during the examination of the Moonabung
Plateau, and to my father, Mr. O. L. Osborne, for his company and assistance on
one trip. Many of the observations on the geology were made in the company
of Professor W. R. Browne, for whose counsel in the field and subsequent helpful
discussion I am very grateful.

Salient Features of the Geology of the Country Surrounding the Area Described.

Before describing the geology of the Lamb’s Valley-Paterson region, it is
necessary to review the geological features of the surrounding country, in order
to have the former area placed in its right geological setting.

K

86 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

On the east and north there is the Paterson-Wallarobba district, part of a
large area of Carboniferous rocks which has been described fully (These
PROCEEDINGS, 1922, pp. 161, 519, and 1925, pp. 67, 112). These rocks comprise both
the Burindi and Kuttung Series, and their structure immediately adjacent to the
present area is anticlinal, being the western side of the Paterson anticline,
so that as one comes west into the area under consideration, higher stratigraphical
units are encountered. On the north-west there are Carboniferous rocks again,
but not very much is known of their structural features. To the west is the
Glendonbrook area and the Stanhope-Tangorin country, both of these being
arbitrarily limited on the east by the line of Lamb’s Valley. This valley runs along
the axis of a northerly-plunging anticline and thus substantially the same section
is found on both sides. To the south-west and south lies the Gosforth-Hillsborough
district; a detailed description of the general geology of this district has been
given by W. R. Browne,’ who shows that the north-east margin of the area
examined by him (the belt bounding the present area on the south-west)
consists of portion of the Lochinvar Dome, dipping north-easterly.

In addition to the Carboniferous areas already mentioned there is an area
of Permian rocks on the south-east of the area considered here. These Permian
rocks have a general basin-like disposition, the lowest member of the Series
marking the boundary of the present map in that neighbourhood.

Thus it is seen that the Carboniferous rocks adjacent to the area have such a
structural arrangement that they dip towards and surround in a general way the
main portion of the area. Thus the chief section of the Paterson-Lamb’s Valley
area constitutes a basin, and this is excellently shown by the plan of the outcrops
on the map. It is proposed to call this the Moonabung Basin, and the plateau
which is the present physiographic expression of that basin, the Moonabung
Plateau. The name “Moonabung” is applied locally as a general district name
for the area. In a former paper (These PRrOocEEDINGS, 1921, p. 261) this was
referred to as Webber’s Creek Plateau.

STRATIGRAPHY AND REGIONAL GEOLOGY.

Preliminary.

The Kuttung rocks occurring at Paterson and to the east were classified by
division into three stages: a Basal, a Volcanic and a Glacial Stage. In the area
considered in this paper, representatives of the last two stages are developed.
Investigations at Gosforth by Browne show that the classification just referred to
also holds for the rocks on the eastern side of the Lochinvar Dome. In the
stratigraphical discussion in this communication most attention will be given to
the consideration of the toscanite-dellenite rocks (Paterson type) and of the
Main Glacial Beds which form the topmost division of the Glacial Stage.

Volcanic Stage.

Rocks of the Volcanic Stage occupy only a minor portion of the district
described. They extend along the right bank of the Paterson River between >
the localities known as Summer Hill and Lennoxton. Just near the latter place
they are displaced by two faults, a small one throwing west and the other, the
important Lennoxton fault, throwing here to the south-east. As a result of the

1Proc. Roy. Soc. N.S.W., 1926, p. 213.

BY G. D. OSBORNE. 87

latter fault the outcrops of the uppermost beds of the Kuttung Series have
suffered a lateral displacement of two miles and a half in a direction perpendicular
to the strike of the beds.

The topmost horizon of the Volcanic Stage is a green albitic rock of the
nature of a pumiceous keratophyre. The upper limit of the stage has been placed
here because the keratophyre is followed by conglomerates and tuffs which form
the Lower Portion of the Glacial Stage. These clastic rocks possess a lithology
which is quite characteristic, and differs from the lithology of the clastic rocks
of the Volcanic Stage.

Two sections may be taken to indicate the variety of units in the Volcanic
Stage. The more important, which is found near Lennoxton, is as follows,
ascending stratigraphically:

Thickness
in feet.

Purple) tuffisiiwathy pebbles. 9-)5 2s 0 ee 4 ee) DO
Toscanite (Mt. Gilmore type) .. .. .. .. 30
Cherts with plant stems a LO Pastene Gc qla Nira tna
Toscanite, albitized (No. 74 5)2 4 Sul Eietne bk ch bti” sts)
Daciter@Nowi43) een ee oe 5, Yaa) Rake ep do.
ACTA Cutis a GINO san 2) mee cere en Une uae an Ba ()
Felspathic tuffs Ss ce teeth, saat Ween) wmosee LOO
Potash rhyolite (No. 741) RON UD ARE DN ec RS 34 yes)
BrecciavViGNoniea nme) Wavy eee eel Ato ge Eh LAE E25)
Felsite (No. 739) .. . Ean SAMI: aes mates O
Green kKeratophyre (No. 746) Bie OM ket aatar poses dite O

Total ase Sere ene 9.0

The second section is found at Summer Hill on the right bank of the Paterson
River near the boundary between the Cessnock and Bolwarra Shires and ascending
stratigraphically, is as follows:

Thickness
in feet.
Bluish¥cure L@No: COG) iinet nie es ecte! esta oO
Coarse banded green tuff (No. 707) .. .. .. 20
Tuffaceous conglomerate (No. 708) .. .. .. 10
Breccia ius ocd ies het isan h cr oe olenrc ane ci
Mauve dacite (No. 713). Be Bone IO
Banded green Keratophyre (No. 709) he apuaten a 1 e0)
MOC Tie 2. eee eee 20D

Some horizons are common to both localities and these can be traced fairly
easily through the intervening country.

The rocks described in the sections just given comprise a series quite compar-
able with the Volcanic Stage sections to be found further east. There is not a
great variety of type, but it should be remembered that in the present case only
part of the Volcanic Stage is dealt with. Of the lavas already detailed, only four
can be correlated directly with units from the districts to the east. These are
those numbered respectively 745, 746, 739 and 741. The particular features of the
more important of the horizons in the Volcanic Stage will be considered under the
heading of Petrography.

1 Numbers refer to specimens in the collection of the Dept. of Geology, University
of Sydney.

88 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

Glacial Stage.
The Lower Portion.

On the eastern and northern sides of the area, this division of the Glacial
Stage (equivalent of the Mt. Johnstone Beds of Sussmilch), consists entirely of
clastic rocks, tuffs predominating. The majority of the tuffs are contaminated
with detrital sediment, becoming in places very pebbly. Conglomerates and sand-
stones are associated with the tuffs, but so far no definitely glacial rocks hav2 been
found, although in this portion of the Glacial Stage to the east, one horizon of
varve-rock has been recorded, viz. from Glenoak, at a position about 200 feet
above the base of the Glacial Stage. On the south-western margin of the area
now being described, however, there are important glacial and fluvio-glacial
deposits. Thus to the north of Hudson’s Peak and Drinan’s Mt., Browne has
shown that the Volcanic Stage is followed by the grey-granite conglomerate and
then a series comprising acid tuffs, varve-rock and tillitic conglomerate, culminating
in a coarse conglomerate consisting of a tuffaceous matrix and many pinx aplite
pebbles. This conglomerate is followed by the Paterson toscanite.

The Paterson toscanite divides the Lower Portion of the Glacial Stage from
the Main Glacial Beds, but between the Moonabung Plateau and Eelah the
toscanite is missing from the sequence, and along this belt it is difficult to draw
the line between the two divisions mentioned above. Still it is quite clear from
what has been said that there is a distinct difference between the rocks under the
Paterson toscanite on the north-east, and their stratigraphical equivalents on the
south-west. This matter is of some interest and importance from the standpoint
of palaegeography and will be discussed under that heading.

Paterson Toscanite and Dellenite.

The rocks which come under this heading immediately underlie the Main
Glacial Beds in all cases except one. In this instance a very thin flow of toscanite
is found sandwiched in with the conglomerate and tuffs which have been described
under the last heading. The occurrence of this was first pointed out to me by
Dr. W. R. Browne, the unit being found on the bridle track from Hillsborough to
Webber’s Creek Falls, just a little to the south of Wildman’s Gap. This
occurrence is of no great importance, stratigraphically, because of its extremely
limited extent.

Concerning the main lava-horizon, it is to be noted that this comprises more
than one flow in the Moonabung area and also to the west of Paterson. At the
base of the main mass on the track to Bell’s house there is a small decomposed
outcrop of pitchstone, seamed with haematite veins. This is identical with a small
flow found in precisely the same stratigraphical position at Westbrook, near
Singleton. Whether this is a glassy phase of the Paterson toscanite, or bears to
the latter a relationship analogous to that shown by the glassy andesites to their
associated lithoidal types is not known.

The maximum thickness of the main mass is about 150 feet, but the varia-
tions in the thickness are very marked. One cannot determine exactly the number
and extent of the subsidiary flows, but it is quite clear that these exist. Thus near
Bell’s house the interposition of conglomerate between two flows of toscanite is
seen. The conglomerate wedges out to the south and the two flows coalesce. In
a number of places a similar state of affairs exists, and in four isolated localities
the presence of two flows without any intercalated sediments has been found. To

BY G. D. OSBORNE. 89

the west of the bridle track not far from the Falls a dense, almost felsitic lava
is found resting on the main flow. This is an albitized rock and recalls many of
the features of the albitized hornblende andesites of Bridgeman.

To the north of Webber’s Creek Falls there are two important flows, which
are both toscanitic in character. In the neighbourhood of Paterson on the northern
side of Webber’s Creek there are a number of interesting varieties of the lavas.
One rock, purplish in colour, shows, under the microscope the presence of a
pumiceous groundmass indicating that it is definitely a volcanic rock. This unit
is associated with two flows of toscanite, which resemble one another.

There are certain features connected with the toscanite in Webber’s Creek
just a little upstream from the Falls, which puzzle one when endeavouring to
interpret their significance. Here there seem to be strings of igneous rock
penetrating the glacial sediments. A similar set of features was seen in the
bed of Dunn’s Creek, east of Paterson and commented upon in 1922. Professor
Browne has remarked upon the features in Webber’s Ck. and suggested that the
toscanite may have been a sub-aqueous flow disturbing the sediments with which
it was associated. From an examination of the toscanites over wide areas during
the past five years I feel fairly sure that they are essentially extrusive rocks,
and that the peculiar features seen in Webber’s and Dunn’s Creeks are probably
best explained by the suggestion of Browne. The hypothesis that the rocks are
sill intrusions raises obstacles that are difficult to remove.

The glaciated pavement discovered in 1921 (see These ProcerepInes, 1921,
p. 259) is upon the surface of the toscanite.

Main Glacial Beds.

Succeeding the Paterson toscanites there is a group of sedimentary rocks
which constitutes the Main Glacial Beds, horizons of glacial origin occurring right
throughout this series. These rocks are characterized by the fact that the
individual units do not persist for any great distance. They outcrop upon the
Moonabung Plateau and have been protected from rapid denudation by reason of
the existence of the basin structure there. On account of the lack of constancy of
the units, it is difficult to find a representative section of any considerable length
from which to obtain a fair indication of the stratigraphical succession. In addition
to this I have not been able to examine the area as fully as desirable, and hence
the stratigraphical detail is not fully known. However, by considering the
succession at a number of places and correlating the data, a good idea of the
general features of the sequence is obtainable.

The rocks immediately on top of the flows of toscanite, etc., vary somewhat,
and comprise varve-rock, tillite, fiuvioglacial conglomerate and tuffaceous con-
glomerate. Careful tracing of the rocks shows that the tuffaceous conglomerates
which are developed at the Bald Hill, north of Bell’s house, are the lowest,
stratigraphically, of the supra-toscanite units. These rocks will be referred to
as the Bald Hill Series, and their maximum thickness is about 350 feet. They are
present to the north of the Bald Hiil, and swing round at the head of Webber’s or
Moonabung Creek, outcropping in Cabbage Tree Brush. They are also found on
the southern slope of Mt. George and above Lennoxton, but they are practically
absent along the stretch of low country to the west of Paterson and also to the
north-east of the glaciated pavement.

The rocks are fairly coarse, there being en abundance of pebbles of quartz

and fine-grained acid rocks like felsites. The pebbles are well-rounded in most
L

90 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

cases and the deposit is reasonably well sorted. Current-bedding is seen in most
places, and the matrix, which is definitely tuffaceous, contains a certain amount of
relatively fresh felspar and biotite. Sometimes the tuffaceous conglomerates pass
into tuffs. The latter type of deposit is found resting on the toscanite to the east
of Mr. M. Boland’s farm. Here there is a section leading up to the plateau which,
starting at the top of the toscanite and proceeding upward, stratigranhically,
is as follows:

Thickness
in feet
Acid tuffs with pebbles eet ids sapere ee megs e100)
Varve-rock bit aaEtE ore aie re ae meek. O)
Conglomerate with current bedding .. .. .. 150
Tuffs with scattered pebbles Dae 40)
Tuffaceous conglomerate 5 RES Be as Mey eee NiO.
Ui Moy oz Nae Pema ise he oe URGXG)

The varves here are somewhat coarse in texture, and they show contem-
poraneous contortions. Further to the east of the point where the above section
is found there is a series commencing with conglomerate in which the average
size of the pebbles is much less than in the rocks of the Bald Hill Series. This
conglomerate contains pebbles of chert, slate, free quartz and felsite. It is
followed by varve-rock which shows a peculiar kind of fracture, whereby large
flaggy pieces become detached from the underlying rock leaving a surface having
a type of confused subconchoidal fracture. This kind of disintegration coupled
with the dip of the rocks has had a definite effect upon the development of minor
ereeks in this region, erosion being facilitated in the direction of dip. Following
the varve-shale are some sandy rocks which may be glacial in origin, and these
are succeeded by tuffaceous conglomerate in which are some bands of tuff. The
pebbles in the conglomerate average about six inches in diameter. The next unit
is a mass of tillitic material containing many angular and subangular fragments
which have been irregularly accumulated. The tillitic material is succeeded by
coarse acid tuff which is ferruginous and varies a little in texture. Then comes
a series of purple shaly rocks reminding one of the Lochinvar shales of the
Permian system, but probably representing glacial shales, similar to rocks of the
same general appearance near the pavement which are definitely glacial. The
shales are followed by coarse conglomerate and this is the highest unit examined
in the section which is summarized as follows:

Thickness
in feet.

Conglomerate aes eae BAN Cuca aetadins hea) Saree Peo lO
WAT VESHMer ch ita 7 crcl tet Be SEE cl ees ee Be ane 80
Sandy, Locks possibly selaciailin sa eis eae 50
AMbuRENeLeYoyots| CLossk=lWopoMNY . G5. oo oo co oo oO)
Tillitic deposit bie Yeas Divan y Wentay . Aeweye cede, ic. ommme OO
Coarse’ ‘acidtufi, «:) 32 “eee eer eet (0.0
Purple shales, possibly glacial .. .. .. .. 150
Goarsescongslomeraten..-) ci) each ann Ee OL

Notal 22. (se. ware elas

If a traverse is taken from the Bald Hill towards the east crossing the basin,
we find that the Bald Hill Series is succeeded by a fine cherty tuff carrying
excellently preserved remains of Rhacopteris and Calamites. This is exposed on
the right bank of Moonabung Creek at the foot of the long spur leading down from

BY G. D. OSBORNE. 91

the Bald Hill. Then comes some very acid tuff, which is immediately followed
by a white plant-bearing tuff. In this Rhacopteris is again abundant throughout
and Calamites stems are limited to thin zones. This horizon is only about 25 feet
thick, and forms a dip slope to the north and north-west of Sherwood’s house,
which leads up to a ridge running to the edge of the plateau overlooking Lamb’s
Valley. The white rock stands out well and forms a useful guide for picking up,
from a distance, the general geographical details of the western side of the basin.

Following the second Rhacopteris-tuff horizon there are some more con-
glomerates, and these are followed by rocks which appear to be the Lochinvar
Shales. These are overlain by Permian conglomerate. On the east side of the
Permian rocks rocks are found which do not fit in with the section on the western
side. Thus there are some varves which are limited to the eastern side, and there
is only one Rhacopteris-bearing horizon. The varves are probably to be placed on
the top of the Bald Hill Series. Varves of the same stratigraphical position occur
at the head of Webber’s Creek in Portion 44, Parish of Wolfingham, resting with
associated tillite on the Paterson type of toscanite.

The Carboniferous rocks in the section just described are as follows:

Thickness
in feet.
Bald Hill conglomerate Bis. vi Aublea lew chic & Wks aa nOR DN)
WEARS thatol TOONS Go loo +906) boo ool oo) bo wll)
JRA DOKGOV OFA abute, IN, U6 5 Bo be 00 100 bo. SH)
Coarse tuff AR Sa Rae aaa eT iota Ree Mis 0
TY CRO GGFES eblit, INOS 7 Go 60 oa 60 Go oo 7)
Conzglomerateeery Gijon, os Doe Wie rte) oreeu en ea00
MOEAIE: (Ra EA ayy:

Considering next the succession in the neighbourhood of the glaciated pave-
ment and Webber’s Creek Falls, we find that immediately on top of the toscanite
at the pavement there are varves with pebbles and a very minor amount of tillite,
while at the Falls the first rocks to succeed the lava are tillites and conglomerates.
Following the varves and tillitic conglomerates come some olive-green to brown
mudstones with scattered plant stems.

Then comes another series of varves which are almost purple or brown in
colour, and relatively free from contemporaneous contortions. These are well
exposed in Bell’s Creek, which runs just to the north of the pavement and joins
Webber’s Creek above the Falls. These varves swing round from this locality and
then die out as we continue across to the northern side of Moonabung Creek; but
they are found on the eastern side of the basin in a number of places. Therefore
this second series of varve-shales is well developed, being fairly widespread in
the area of the basin.

A little north of Bell’s house we see an occurrence of varve-rock which,
as regards stratigraphical position, is of interest. 'The glacial rock is associated
with conglomerate which can be followed down the small creek just to the north
of Bell’s. This conglomerate clearly separates two flows of toscanite and hence the
varve-shales are sandwiched in between the lava horizons. From the genera! field
evidence I think it very probable that these varve-shales are lower stratigraphically
than those at the glaciated pavement and in Portion 44.

The stratigraphy of the rocks on the Moonabung Plateau, ascending strati-
graphically, is summarized as follows:

92 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

Carboniferous.
Thickness
in feet.
(Maximum
values. )
Main toscanite unit Nee ha hepa bis ig quam lSXt)
Varve rock and conglomerate a Bell’ Sioa seas 90
Subsidiary flows of toscanite, cte. .. .. .. 100
Bald Hill Series .. . ieeaeye © 3.0.0
Pavement varve-rock and tillitic deposit sg abaxt)
Mudstone with plant stems s eaahece ee ee Se 90
Rhacopteris tuff No. 1 Syd fa Scy pagans, ey ee anes 30
Acid, ferruginous tuffs Buel Sure ke Dre ee 80
Rhacopteris tuff No. 2 Ch BW WC 3 A ee 25
Purple varve-rock Brel aneeD eth ely Le eel 0
Conglomerate Ba Near ie alien at Usa ae ami Dey)
Coarse acid tuff ... slye se st bees enero LOO)
Purple shales, possibly elesien Pee Nee e amris Sy eben oy GEER)
Coarserconslonierates ape eet ene 70
TPO tai ae em apo
Permian.
Shales (probably the Lochinvar Shales) 5% 50
Amygdaloidal basalt .. .. oi 20
Tuffaceous sandstones and conclameratas sa. PAY)
TRO EALMIE ea trses Secreta GO

This -gives a thickness of 1,475 feet for the Main Glacial Beds, which is
distinctly less than that obtained at Seaham, but many of the values for thickness
given here are only approximate.

It is necessary now to consider the rocks west of Paterson and between
Lennoxton and the Permian area on the south. Along the left bank of Webber’s
Creek for some miles west of Paterson it is possible to gather some information
about the succession. Just near the Paterson Park there are two bands of varve-
rock with associated sediments, and one can generally find evidence of the
existence of the same two horizons as one goes west. Associated strata comprise
fine white tuffs, coarse red tuffs and coarse conglomerates with tillitic phases, in
one of which an erratic about five feet in diameter was found. The second set of
varves thickens considerably to the west and is found well developed towards
the Falls, attaining to a maximum of 150 feet.

As one goes up Smith’s Creek to the north one finds definite tillite along with
the varve rock, and to the south of Lennoxion above the Paterson River the
following section in the Main Glacial Beds is found:

Thickness
: in feet.
Conglomerate, probably equivalent of Bald
Hill Series Ae MP8 $ ae Sap EE 90
Wellow. Shute) seme + cal eh a ORS 80
Coarse tuffaceous UBIO. SO ERED oo ANAKD
Tillitic deposit ye : 20

Normal conglomerate, Rvith! pine mpiite Henoles 80

MoOcalumerer a ion e AL

It will be seen that the varve-rocks have disappeared from this section.

BY G. D. OSBORNE. 93

Proceeding from the Falls towards Rosebrook the varves are seen to be
strongly developed, the horizon being the equivalent of the purple varves on the
Moonabung Plateau. On the eastern fall of the Rosebrook ridge and to the north
of Drinan’s Mt. there are, of course, at least two horizons of varve-rock.

Associated with the glacial beds just east of the Falls there are some
siliceous tuffs which are not seen elsewhere.

Outliers of Permian Age.

Apart from the large area of Permian rocks in the south-east of the area,
there are a number of outliers. Most of these are only a few acres in extent,
but a larger one is to be found to the east of the Falls (see map). It is probable
that there exist other small outliers which are not shown on the map. The rocks
constituting these outliers comprise units which are sometimes representatives
solely of the Lower Marine Series; as for example the Lochinvar Shales and
the amygdaloidal basalt. At other times there are conglomerates which remind
one of the Upper Marine conglomerates south of Paterson.

The height above sea-level of the base of the outliers on the Moonabung
Plateau is fairly constant, and is in excess of the height of the same horizon on
the east side of the Falls by about 340 feet. These data confirm the existence of
the Lennoxton fault and to some extent give a measure of the throw in this
locality.

STRUCTURAL GEOLOGY.

The greater part of the area consists, structurally, of a basin which has been
broken by a large fault on its eastern side. The examination of the dips shows
the existence of this basin quite clearly, and physiographically the structure
stands out well, as the toscanites, particularly the main one, have on account of
being resistant units, dominated the topography. The section given in Text-figure 1
shows the structure of the Moonabung area. The Moonabung basin is of the
same general nature as the other basins in the district to the west, viz. the Cranky
Corner and Mirannie Basins, and it is separated from the latter by a comple-
mentary structural feature. This is a plunging anticline, out of which Lamb’s
Valley has been carved. The anticline is really the northern end of the Lochinvar
Dome, the northerly dip of the beds being the measure of the pitch of this portion
of the dome.

Just as the Lochinvar Dome is elongated in a more or less meridional direction,
so the Moonabung Basin shows an elongation in that direction, and the plan of the
outcrop of the rim of the basin (which consists chiefly of toscanite), is an ellipse,
the length of the major axis of which is about five miles, and of the minor axis
not more than three miles.

The eastern portion of the area, which is separated from the Moonabung
Plateau by the Lennoxton Fault, forms the western side of the Paterson anticline.
This structure was described in a former paner in 1922, although much of the
detail of this portion of the region was not known then. It will be seen by
inspection of the dips to the west of Paterson that the toscanite is dipping in a
general southerly direction for some distance, and there is a change in the dip
further west, the beds becoming flat near Lennoxton. The structure in the
last-named locality is shown by considering Text-figure 2 in conjunction with
the map. The gentle dip of some of the beds and the advanced state of the
dissection have brought about the existence of both outliers and inliers of certain
horizons.

S VALLEY

GEOLOGY OF PATERSON-LAMB

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BY G. D. OSBORNE. , 95

In addition to the Moonabung Basin and the eastern region near Mt. Johnstone,
both of which have been already considered, there is a third region which comes
up for consideration. This is the area in the south-east which is given over to
Permian rocks. The detail of the structure of this area is not yet known and we
can only refer to the general structural features. The junction of the Permian
and Carboniferous systems from Eelah to the north can be followed fairly readily
as has been shown by W. R. Browne, by noting the outcrop of the Lochinvar
shales. These appear, however, to cut out as one goes north from the neighbour-
hood of Rosebrook, except that they are found in the Permian outliers in a number
of places. However, they are definitely missing from the sequence near Paterson.
A consideration of the strata in the field has led to the position of the junction of
the two series being placed as shown on the map.

Now it will be seen that the Permian rocks lie in a plunging syncline, or, to
borrow a term used by Prof. Browne, form a “gulf” in the Carboniferous area.
This syncline is complementary to the Paterson anticline on the north-east side,
and to the eastern margin (now faulted) of the Lochinvar Dome, on the south-
western side. But associated with these relationships we have the rather
abrupt change in the dip direction of the strata in the neighbourhood of the S.W.
end of the Lennoxton fault. Hereabouts one finds on the S.E. side of the
postulated position of the fault, the beds turning round to strike in sympathy
with the Permian syncline. On the N.W. side of the fault line, however, the beds
dip in accordance with the Moonabung Basin. These facts emphasize the irregular
nature of the folding.

Faulting.

The most important fault in the area is the Lennoxton fault. This has been
followed from a point a little to the south of Wildman’s Gap to Vacy on the
Paterson-Gresford road. The fault is clearly shown by the outcrops of the various
horizons, and it is manifested physiographically also. Above the river at Lennox-
ton there is a parallel fault associated with the larger one, the former throwing
to the west. From a consideration of the structure to the east of Webber’s Creek
Falls it is clear that the Lennoxton fault is normal in character, throwing in
general to the east and south-east. The throw of the fault is difficult to estimate,
and it increases as one goes north, but in the neighbourhood of Lennoxton it is
between 2,000 and 2,500 feet.

It is rather singular that this dislocation is absent along the track from
Hudson’s Peak district to the Falls. On the S.W. side of the track at Wildman’s
Gap there is an interesting fault which has been described by Browne. This and
the Lennoxton fault are almost collinear, but careful consideration of the problem
and detailed field examination seem to prove clearly that the two are unconnected.
The fault in Kilfoyle’s creek is not a large one and may easily die out north-
eastwards, but the other dislocation seems to be of fair magnitude near the Falls,
and one would expect to find it crossing the track to Wildman’s Gap. But this it
does not do, and it can only be surmised that the fault does not extend any further
than is shown on the map. Just at the limit of the map in Portion 9 Parish of
Wolfingham there is a number of small outcrops of the Paterson toscanite, and
these seem to be the displaced equivalents of the main mass outcropping on the
Moonabung Basin. They are of a lower altitude and an examination of their
general features of occurrence suggests that they are the remnants of the toscanite

96 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

flows, which thin out very definitely as one comes S.H. from the Moonabung
Plateau, and that here we have a small displacement along the fracture of the
Lennoxton fault, the amount of displacement increasing as one goes north.

Beyond Lennoxton the fault turns to the east, and near Vacy is hidden by thick
alluvium, but it looks as if it carries on and becomes the fault described as F, and
shown on the Clarencetown-Paterson map (see These PROCEEDINGS, 1922,
Plate xxvi), since the latter possesses the same features of strike, magnitude, etc.

A very striking effect of the fault is the displacement of the higher lavas of
the Volcanic Stage from their alignment at the foot of Mt. Johnstone to the right
bank of the Paterson River near Lennoxton.

In addition to the large fault just described, there are a number of smaller
ones. One already mentioned throws to the west. Another occurs on the east
side of Lamb’s Valley above Mr. Boland’s farm, where the displacement of the
toscanite is clearly seen. Further, in many places on the Moonabung Plateau the
beds are standing on edge, indicating faulting, although very few details can be
obtained regarding the nature of the movements. Frequently also on the Moona-
bung Plateau strong jointing is observed, and this feature is no doubt the result
of faulting movements.

Finally near Paterson along the Webber’s Creek road there are small faults
displacing the junction line between the Permian and Carboniferous Series.

All the faults are normal and belong to a general group which includes those
described from the Paterson-Seaham region and the series of normal faults mapped
in the Singleton district. All these normal faults are probably of late-Palaeozoic
age for reasons given in two earlier papers (These PRocEEDINGS, 1922, p. 525, and
1926, p. 401).

PHYSIOGRAPHY.

The area being considered is part of the northern margin of the valley of the
Lower Hunter River. Dissection has proceeded to a fairly advanced degree, and
the aspect of the topography on the whole is that of late-maturity, although there
are exceptions where some of the smaller creeks are concerned, and also some-
times the same creek will show both youthful and mature aspects in different parts
of its extent. Thus in the case of Webber’s Creek we find an interesting assemblage
of varying physiographic features as we pass upstream.

The physiography of the area is intimately bound up with the geological
structure, and thus it is convenient to maintain the division used in the chapter
on the structural geology, taking the Moonabung Plateau as one part and that
constituting the remainder of the region as the other.

The faulting which has occurred does not manifest itself in the presence of
initial fault scarps, because the fault-scarp-formation antedated the peneplanation
and subsequent uplift, which were in turn antecedent to the present cycle of
erosion.

However, the manner in which the strata were arranged by the Lennoxton
fault coupled with the arrangement due to folding has had a marked influence
upon the topographic evolution.

The peneplain surface in the area of the map was probably not so regular
as was the case in some parts of the adjacent country, as for example from Mt.
Johnstone towards Mt. Gilmore. What are probably remnants of some of the
original shallow valleys and rounded hills are to be seen in certain parts of the
Moonabung Plateau, it being fairly clear that Mt. Johnstone and Mt. George and

BY G. D. OSBORNE. , 97

the Bald Hill as seen now have not suffered much lowering of height due to the
erosion which followed the uplift. The respective heights of these, viz. 1,130, 1,466
and 1,050 feet give some idea of the height of the plateau in this region at the
completion of the Kosciusko uplift.

In two or three places one can see examples of valley-in-valley structure,
generally on a small scale, the original broad valleys developed in late Tertiary
time, being cut into by small streams, which in these cases are the rejuvenated
representatives of the older streams.

The adjacent area of the Paterson-Clarencetown district and the area under
consideration belong to the same physiographic unit (part of an uplifted
peneplain), and there is as one goes from the west towards the south-east from
the present area, a decrease in the general height of the plateau, this probably
being due, as mentioned by Browne in the paper on the Gosforth district, to an
original slope in the surface of the plateau when uplifted.

There are a number of outliers on the surface of the Moonabung Plateau.
These are remnants of the Permian rocks which at one time covered much of the
area now forming the surface of the plateau. These rocks were at one time
continuous with the Permian rocks south and west of the Hunter River at
Gosforth. The denudation of much of the capping of Permian rocks was accom-
plished in pre-Kosciusko times, and thus the uplifted peneplain was characterized
by the existence of relatively isolated patches of Permian rocks scattered about
on the Carboniferous basement, these being subsequently somewhat reduced in
size by Pleistocene erosion.

As the physiography is affected by the nature of the rocks it is interesting
to note the varying resistance offered by the different units to erosion. The most
resistant rocks are the toscanite and dellenite (Paterson type). These are respon-
sible for many striking dip slope-escarpment elements which are wholly or partly
made of these rocks. The toscanites of the Moonabung Plateau produce the ring-
like outcrop which bounds the plateau along almost the whole of its extent. The
precipitous cliffs on both sides of Lamb’s Valley, which are inaccessible in places,
are made of the toscanite.

The conglomerates in the Lower Portion of the Glacial Stage stand out much
more strongly than the tuffs and varve-rocks. A good illustration of this is to be
seen on the bridle-track to the Falls. Here one crosses two bands of coarse
conglomerate separated by tuff, and the former produce steep “pinches” on the
track while the tuff has been eroded rather rapidly from off the surface of the
lower conglomerate thereby allowing for the presence of a gently sloping section of
track along to the base of the upper mass of conglomerate.

The Main Glacial Beds as a whole offer only a medium degree of resistance
to denudation, the Bald Hill Series being the exception. The rocks of this series
are fairly well cemented conglomerates and tuffs and they are generally found
standing out forming fairly rugged features as at the Bald Hill. The varves and
mudstones break up rather easily and the tillitic conglomerates, on the whole,
disintegrate readily.

In the area of the Volcanic Stage rocks it will be noticed that a broad valley
has been eroded by the Paterson River, but while this is a reflection of the
advanced state of the denudation, it also emphasizes the fact that the lavas of
that stage, on account of their lack of any great thickness in all cases, have not

resisted erosion and along with the easily disintegrated conglomerates have been
rapidly removed.
M

98 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

The most important stream in the area is the Paterson River, but we are
only concerned with the extreme margin of its valley in this area. ‘The
physiography of the Lower Paterson has already been described by the writer in
another paper, but it is interesting to record the presence near Lennoxton and
Summer Hill of terracing along the Paterson. The exact cause of the production
of these is not known, and while they may indicate the occurrence of small uplifts
late in the history of the river, they may be due to uninterrupted erosional
activity.

Dismissing, then, the Paterson River, we see that the next most important
stream in the area is Webber’s or Moonabung Creek,’ generally called by the latter
name in its upper portion. This stream joins the Paterson River some two and
a half miles south of Paterson and its valley for some distance upstream is heavily
alluviated. From the point of confluence to the west for a distance of about four
miles, the creek is flowing in a direction more or less parallel to the strike of the
rocks, and because of this it has been able to erode a fairly wide valley along
this belt. The northern limit of the valley just west of Paterson is the outcrop
of the Paterson toscanite, and to the south the boundary is, in part, made by the
sandstones and conglomerates of the Upper and Lower Marine Series. Thus the
Main Glacial Beds and the Lower Marine cherts, etc., have been extensively
removed.

Further along the valley, one finds that the trend of the creek and the direction
of strike of the rocks are no longer more or less parallel. Eventually, one reaches
the barrier of toscanite at Webber’s Creek Falls, where, due to the irregular
folding and the Lennoxton fault, we find the creek flowing directly across the
strike. The falls are about 90 feet in vertical range and form an imposing sight
in winter time immediately after rain. They are carved out of toscanite almost
entirely.

Above the Falls, Webber’s Creek presents a valley, different altogether from
that seen on the lowland to the east of the Falls. Now we find that the stream
and its tributaries are not able to erode quite so rapidly and on account of the
centroclinal nature of the dips on the plateau there is not any striking widening
of the small valleys in any particular direction. The presence of the barrier of
toscanite at the Falls has brought about the deposition of alluvium over a relatively
wide area upstream, and this has been contributed to by the direction of the dip
of the strata just above the Falls being directly opposed to the direction of fiow
of the creek. The alluvium so accumulated produces swamp-like areas in the wet
seasons and the aggradation has prevented, in some measure, the deepening of the
channel of Webber’s Creek to the north-west of the Falls.

The tributaries on the north side of Webber’s Creek in the region to the east
of the Falls have developed much more extensively than those on the south, and
this is due, partly, to the southerly-directed dip of the rocks in the major portion
of the northern area. Thus Smith’s Creek, and the two to the east of it, have
fairly broad valleys, and the extensive erosion has brought about the existence of
both inliers and outliers of certain units.

Professor Browne, in discussing the physiography of the Gosforth district, has
suggested that in the evolution of the present Hunter River which began with the
epeirogenic movements at the close of the Tertiary, there were two main stages of

1[t should be noted that there is another Webber’s Creek, viz. in the Glendonbrook
district.

BY G. D. OSBORNE. 99

erosion consequent upon two uplifts. The former stage consisted of denudation of
Permian sediments and the production of certain river and creek systems, which
were superimposed upon the underlying Carboniferous terrane when the second
uplift took place.

With regard to the physiography of the Moonabung area, as a whole there is
nothing to show whether or not there were two main stages in the late Tertiary
uplift, but there is evidence of rejuvenation of the ancient streams having
occurred.

PALAEOGEOGRAPHY.

The palaeogeography and geological history of the Gosforth district and its
environs have been ably dealt with by Dr. Browne, and much of what he has
written will apply to the Moonabung area because this is intimately related
geologically to the former region. However, there are one or two points connected
with the present district which may be considered here: The extent of the Paterson
lavas and the nature of the sediments in the Lower Portion of the Glacial Stage
and their probable former extent in the region being discussed.

It is quite clear that the toscanite and delienite flows were very widespread
when one remembers that they have been found from Raymond Terrace to at least
as far as Mt. Dyrring, north of Singleton. On the north their limitation is not yet
known, but while they are found close to the base of the Permian along from
Seaham towards Moonabung, they are definitely missing from the sections east of
Rosebrook and near Helah, and probably they do not occur in the Gosforth region,
although there are some toscanite flows in the glacial beds recorded by Browne
for the west of the Hillsborough district.

The remnants of the thin margins of the toscanites are to be seen between
Drinan’s Mt. and Wildman’s Gap and the flows on the Plateau above the Gap thin
out east and south. That the thinning is rather abrupt is suggested by the fact
that at the Falls the lavas are about 90 feet thick while a little to the east on the
other side of the Lennoxton fault there is no sign of the igneous rock in the
succession. Thus in short at a time just prior to the accumulation of the Main
Glacial Beds there developed the interesting group of toscanites and dellenites,
the flows spreading over a vast area covering the underlying sediments to a depth
of about three hundred feet at Paterson where we find the greatest development of
the rocks. The area occupied by the igneous rocks was bounded on the south
(in this particular region) by a line running from near Wildman’s Gap towards
the east.

With regard to the rocks constituting the Lower Portion of the Glacial Stage
one notices the constancy of the basal conglomerate, but for the bulk of the
remainder there is a distinct difference in facies between the rocks which are
found right throughout the Clarencetown-Seaham district and to the north of
Moonabung on to Mt. Dyrring, and their stratigraphical equivalents in the region
extending from just north of Drinan’s Mt. to Eelah, and also in the Gosforth
district. The rocks of the former areas consist of tuffs with subordinate con-
glomerates which occur in bands. The congiomerates are not coarse-grained, as a
rule, and are always well cemented. The tuffs possess a brown, sometimes
chocolate, colour and have a lithology which is characteristic. The rocks of the
second group are characterized by the prevalence of pink-granite conglomerates
and the general lithology is just as typical as, and quite distinct from, that of the
other series. One can see the intermingling of the two types of sediment as one

100 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

goes from north of Drinan’s Mt. along the base of the Plateau. The pink-granite
type of conglomerate is seen on the track to Bell’s, but it loses its individuality
as we proceed to the north, and at the head of Lamb’s Valley and on the northern
slope of Mt. George we have the tuffs and more finely-textured conglomerates of
the group which is so well developed in the Paterson region. Further, at Gosforth,
the rocks of the Main Glacial Beds comprise pink-granite conglomerates, but such
sediments are practically absent from the Moonabung Plateau, although imme-
diately following the Paterson toscanite there are conglomerates (the Bald Hill
Series) which display the general facies of the Paterson-Singleton group.

Browne has pointed out the probability that the area, which provided the
granitic pebbles, which are so prevalent in the conglomerates right through the
Kuttung Series in the Gosforth district, was situated to the south, and with this
I am in accord, but from a consideration of the data to hand regarding the distri-
bution of the two types of sediment, which are found in the Lower Portion of the
Glacial Stage through the areas to the north of Gosforth, I would suggest that
the following course of events characterized the development of the Lower Portion
of the Glacial Stage. While the conglomerates and other sediments of the present
Gosforth district were accumulating in post-Voleanic Stage time in the southern
portion of the large Upper Kuttung lake, volcanic activity was manifesting itself
by ejecting fragmental material which was widely spread. The material falling
in the Gosforth district, was, taken as a whole, distinctly coarser, and less
contaminated with detrital sediment, than that which accumulated throughout the
large area from west of Singleton to Clarencetown. Over this latter region there
was being piled up a series of tuffaceous sandstones and occasional conglomerate,
the latter indicating grand flood periods in the cycie of erosion. The nature of
the sediments in this region and the presence of current bedding suggest that
these were developed in a large delta. It is probable that this delta in the great
Kuttung freshwater lake, flanked the north-western shore, the areas providing
the detritus for the delta formation being composed of a terrane, different from
that of the area to the south, where granitic rocks abounded. A line which would
effect a broad division between the two types of sediments in the Lower Portion
of the Glacial Stage could be drawn on the map, and the actual physical boundary
would pass under the Moonabung Plateau.

PETROGRAPHY.

The petrography does not need to be fully discussed here, because the general
petrological features of the rocks correspond very closely with those of the rocks
of the Clarencetown-Paterson region, which have been described in an earlier
paper (These ProceEepincs, 1925, pp. 112-138). There it is pointed out that, in the
region mentioned, there is a great variety of lavas ranging from intermediate to
acid types, and embracing andesites, dacites, keratophyres, toscanites, dellenites,
rhyolites and felsites. Some of these are vitrophyric, and some, particularly the
more acid, are tuffaceous. Associated with the lavas are some acid albitic tuffs
and flow breccias.

In connection with these rocks it is also shown that on account of albitization
having affected them it is difficult to make out the magmatic relationships and
to work out problems of petrogenesis. On the other hand, it is possible to recognize
only a broad sequence of flows throughout the area. Amongst the salient petro-
logical features shown by these rocks, when considered as a whole, are the

BY G. D. OSBORNE. 101

following: evidence of the operation of late magmatic processes effecting albitiza-
tion and kaolinization; autobrecciation; devitrification; spherulitic structure.

Coming now to the rocks of the Moonabung-Paterson region we note that
many of the rocks correspond in general features to rocks from the more eastern
district. The sequence which is found in the upper part of the Volcanic Stage
near Lennoxton is as follows: Toscanite, albitized toscanite, dacite, potash rhyolite
and keratophyre.

Specimen 745 is a toscanite from Lennoxton, which in hand specimen shows
a general brown colour with lighter streaks due to the presence of elongated
inclusions. Free quartz is very abundant and a certain amount of felspar is
visible. Microscopically, one sees much phenocrystic quartz and some andesine
which has suffered albitization, together with a little orthoclase and some biotite.

These are set in a crypto-crystalline groundmass which exhibits flow structure
rather well.

Specimen 743 from Lennoxton is a dacite which consists of phenocrysts of
quartz, which are often well crystallized, and andesine, set in a rough-feeling
groundmass which is a devitrified pumice.

The next flow after 743 is a potash rhyolite of which 741 is a specimen. This
is always of a red colour due to the presence of haematite which has developed by
devitrification of a glassy rock. The haematite occurs in patches and the structure
of the pumiceous groundmass is well displayed by the outline of the iron oxide
patches. The felspars are clear and not albitized, and in addition to fairly
abundant orthoclase there is some oligoclase.

Specimen 740 is from the felsite at Lennoxton which is near the top of the
Voleanic Stage. The rock is light in colour, and has the rough feel characteristic
of trachytes. In thin section it is seen to be almost entirely a devitrified pumice,
in which are set phenocrysts of corroded quartz, albite (which is probably of
deuteric origin), and subordinate biotite.

Specimens 710 and 713 are from near the Paterson River at Summer Hill,
being just at the top of the Volcanic Stage. These are phases of the same lava,
the general nature of the rocks being dacitic. Some of the felspar is completely
albitized but remnants of original andesine can be made out. The horizon from
which these examples were taken is characterized by the variation which is found
along the strike.

The green keratophyre which, as shown in the stratigraphy section, is
generally at the top of the Volcanic Stage, possesses fairly constant petrographic
characters, although there is a great variation in the colour of the rocks from
place to place. In hand specimen the white felspar phenocrysts stand out well,
and these are seen under the microscope to be albite. The groundmass is strongly
pumiceous, and at times there are examples of autobrecciation. These rocks may
be called tuffaceous soda rhyolites or keratophyres.

As examples of the tuffs of the Volcanic Stage we may take Specimens 706 and
742. The former is from near Summer Hill on the Paterson River, and is a
dark-blue rock which is really a flow-breccia, consisting of fragments of felspar and
quartz, and also of dacitic rocks included in a pumiceous groundmass or matrix
which in places shows flow structure.

No. 742 is a quartz tuff which consists of quartz, albite orthoclase and biotite
cemented by haematite and kaolin. In addition to the mineral fragments there
are occasionally angular pieces of cherty rocks.

102 GEOLOGY OF PATERSON-LAMB’S VALLEY DISTRICT.

While considering the tuffs of the Volcanic Stage, brief mention can be made
of the tuffs which make up such a large proportion of the Lower Portion of the
Glacial Stage. Specimens 705 and 712 are typical of these tuffs, both being from
the Summer Hill locality. These are similar in general features, although differing
in detail, the former, for example, containing much more in the way of rock
fragments. Both rocks consist essentially of pieces of quartz, and plagioclase
(often showing albitization), and fragments of glassy and cryptocrystalline rocks.

Paterson toscanite and associated lavas.

As was the case in the other areas, it is convenient to treat the rocks embraced
under the above heading as a separate group. In addition to the toscanite, and
the closely allied dellenite, we find in association, especially where there is more
than one flow, soda rhyolites and certain slightly tuffaceous rocks. Probably the
most important feature of this group for the present area, in comparison with the
same group in the Paterson area, is the occurrence below the main mass of
toscanite of a pitchstone, which shows a certain amount of devitrification.
Specimen 730 of this unit is of a reddish colour and possesses an irregular
fracture. Phenocrysts of felspar and biotite can be seen quite clearly in hand
specimen. Under the microscope the phenocrysts comprise quartz, andesine, biotite
and magnetite. The first of these is somewhat angular in piaces, but generally
corroded. The andesine varies in grainsize but maintains a tabular habit. It is
fairly fresh and any alteration seems to be the result of ordinary weathering. 'The
mica is notably altered, being bleached and strongly resorbed at times, and the
majority of sections show a pleochroism from golden yellow to deep brown.
The groundmass is strongly haematitic due to devitrification, but the greater part
is still glassy, and exhibits flow structure. The absence of pyroxene and
amphibole phenocrysts is of interest and suggests that the rock is distinct from the
pitchstones in the lower part of the Volcanic Stage. The rock also differs from
many of the rhyolitic pitchstones which are found near Glenoak, and which
contain no visible biotite. A chemical analysis is needed to discuss the relation-
ship of this rock to the associated toscanites, but it is probable that a fair amount
of orthoclase exists in the base.

Specimens 259 and 260 are from a spot about three miles west of Paterson
where there seem to be at least two flows in association. The former is a
toscanite, possessing a fawn colour, while the latter is a dellenite and possesses
a dark blue colour. Wo. 259, when examined microscopically, shows individuals
of quartz, orthoclase and strongly albitized plagioclase with occasional biotite, set
in a groundmass which appears to have been devitrified. No. 260 differs a good
deal from the former and shows the presence of much biotite with quartz and
the two felspars, in a groundmass which exhibits excellent examples of fluidal
fabric, and which possesses some pumiceous patches and a little tuffaceous material.
There is no doubt of these rocks being of extrusive origin, and hence the
importance of this evidence in relation to the problem of the mode of occurrence
of the Paterson toscanite suite.

Near Bell’s and elsewhere there are definite cases of the existence of two
flows. Specimen 737 is from the upper flow just north-east of Bell’s house. It
differs from the main mass in that there is more phenocrystic felspar and less
quartz. The general features of the groundmass are similar in both cases.

Amongst the sedimentary rocks (excluding primary tuffs) there are many
iuffaceous sandstones. No sections of these have been made, but macroscopic

BY G. D. OSBORNE. 103
examination gives some information about their constitution. Generally there
are grains of quartz and felspar and tiny chips of fine-grained rocks cemented by
argillaceous material, with a minor amount cf haematite. Sometimes the haematite
is strongly developed and characteristic red rocks result. Then again, in a few
isolated instances there are examples of rocks which possess a siliceous cement.

Concerning the conglomerates one has to note that the matrix is often
arkosic, and may have been derived entirely from detritus, or more generally may
contain a certain amount of fragmental material derived from volcanic centres.
In the case of the Bald Hill Series, biotite is quite an important constituent.

The general nature of the pebbles of the conglomerates has been indicated in
the discussion of the stratigraphy and palaeogeography.

EXPLANATION OF PLATE I.
Geological Map of the Lamb’s Valley-Paterson District.

A NEW DELTOPECTEN FROM THE ILLAWARRA DISTRICT, N.S.W.

By Joun MircHe tt, late Principal of the Technical College
and School of Mines, Newcastle, N.S.W.

(Plate ii.)
[Read 30th March, 1927.]

DELTOPECTEN RIENITSI, n. sp. Plate ii, figs. 1 and 2.

Spec. chars. Equivalvular, very inequilateral, suboblique, outline suboblong.
Right valve strongly convex, ears apparently subequal, transversed by numerous
radii, lati-transverse folds of the anterior ear strongly arcuate, and distinct finer
striae numerous; radials of the body fine, increased by interpolations, and before
the ventral margin is reached are grouped into distinct fasciculi, consisting of six
to ten radii in each bundle, the transverse striae are fine and echinate the radials
where these are crossed by them; the wider growth lines have scalloped or wavy
edges. Beak prominent, incurved, pointed and reaching beyond the hinge line.
Left valve flattish, radials less numerous and convex and less distinctly fasciculated
than those of the right valve; the beak, too, is much less prominent than is that
of the right valve; the two valves are distinctly agape along the hinge line,
resilium large. The anterior ear protrudes beyond the anterior end of the shell,
and is strongly emarginate.

Dimensions: Length 7:2, width (depth) 4:5 cm.

This Deltopecten resembles D. obliquatus Ethr. and Dun. (Mem. Geol. Survey,
N. S. Wales. Pal. No.5, 1906) in several features but it differs from that species
in having a large resilium, fasciculated radials, more apparent on the right valve
than on the left; and less oblique in shape.

Loc.—lllawarra District.

Horizon.—Upper Marine beds. Permocarboniferous.

Dedicated to Mr. H. G. Rienits, of Mt. Victoria, N.S.W., who allowed me the
privilege of describing this fine specimen and who also kindly presented the type
to me.

THE FOSSIL ESTHERIAE OF AUSTRALIA. Part i.

By JoHN MitTcHELL, late Principal of the Technical College
and School of Mines, Newcastle, N.S.W.

(Plates ii-iv.)
[Read 30th March, 1927.]

Introduction.

The earliest record (Cox, These PROCEEDINGS, 1880, p. 276) of the occurrence
of fossil Estheria in Australia was made by Dr. J. C. Cox from the Moore Park
Diamond Drill bore; and to this first specimen Dr. Cox gave the name ZFstheria
coghlani, after Mr. John Coghlan of the Diamond Drill Company, who was directing
the work of sinking the bore referred to, but he neither described nor figured the
species. The species was subsequently described by the late R. Etheridge, Junr.
(Mem. Geol. Survey, N. S. Wales, 1888, Pal. No. I, pp. 6-8, Pl. i, figs. 1-10). The
next reference to the occurrence of fossil Estheriae in Australian rocks was also
made by R. Etheridge Junr., who in 1892 referred some specimens of this genus,
obtained from Denmark Hill, near Ipswich, Queensland, to HE. mangaliensis Jones
(Jack and Etheridge, Geol. and Pal. Queensland and N.G., 1892, p. 387). In 1909
I recorded the discovery of Estheriae in the Newcastle coal measures (Notes and
Exhibits, These Procreprnes, 1909, p. 411-12).

In 1925 I referred to the presence of Estheriae in the Belmont chert (tuff)
beds. (Descriptions of new species of Leaia, These PROCEEDINGS, 1925, p. 438.)

During the year 1890 I collected several specimens of HEstheriae from the
Wianamatta series near Glenlee, but did not record the find at the time.

It will be seen by the above record that the geologic range of Estheria has
been considerably extended since 1892, which will be obvious from the following
list of the series of rocks from which they have been collected to date, viz.:

1. Wianamatta Series, about 700 feet thick (Clarke) (but probably exceed 1,009 feet
in the vicinity of Cobbitty). ‘

2. Estheria Shales, about 610 feet thick (David).

8. Newcastle Coal Measures, 1,100 or more feet thick (David).

To complete the actual stratigraphical range of these fossils, the intervening

series between the Wianamatta and the Newcastle Coal Measure must be included,
as: follows:

1. Wianamatta series, about 700 feet thick (Clarke).

2. Hawkesbury Sandstone, about 1,000 feet thick (Wilkinson).
3. Narrabeen Shales, about 650 feet thick (David).

4. Estheria Shales, about 610 feet thick (David).

5. Newcastle Coal Measures, about 1,100 feet thick (David).

The total thickness of these together exceeds 4,000 feet; but that does not by
any means represent the full geologic range of these fossils in New South Wales

as far as present knowledge discloses, for there are at least two gaps in the portion
N

106 FOSSIL ESTHERIAE OF AUSTRALIA, i.

of the geologic record shown above, and these represent a considerable length of
time even from a geologic point of view. These gaps occur between the
Wianamatta beds on the one hand, and the Hawkesbury Sandstones on the other,
and again, between the Hstheria Shales and the Newcastle Coal Measures.

That the gap between the close of the Hawkesbury Sandstone Series and the
commencement of the deposition of the Wianamatta Series thereon represents a
considerable length of time is shown by the extensive weathering the former had
undergone before conditions arose to admit of the formation of the latter; and that
the break between the Newcastle Measures and the subsequent deposition upon
them of the Mesozoic measures represents a great length of time, is shown by the
great changes in the fossil Flora ushered in with the Mesozoic deposits lying
immediately upon the Newcastle measures. The length of this lapse of time is
further emphasized when the denudation of the lower measures before une upper
series began to be laid upon them is also considered.

The Mode of their Occurrence.

In the lower portion of the Hstheria Shales and in the Wianamatta Series they
occur in thin bands of impure clay ironstone. In-the Newcastle Measures at
Belmont and at Warner’s Bay, they are found in cherts (tuffs). Judging by the
fossils associated with them, it would appear that in each case they were dwellers
in fresh, or slightly brackish waters. Those from Belmont and Warner’s Bay occur
as casts only; but those from Merewether Beach, Newcastle, from the Estheria
Shales and the Wianamatta beds often have tests preserved. Those obtained from
the Estheria Shales and the Wianamatta Beds are black and lustrous, but those
from the Newcastle beds (Merewether Beach) are of a dun colour. It is not
uncommon to find specimens with two valves conjoined, but never free from the
matrix, so far as I am aware.

The variety of forms of Hstheria obtained from the Newcastle Coal Measures
is remarkable.

From Victoria, South Australia, West Australia and Tasmania, the occurrence
of fossil Estheria, so far as I am aware, remains to be recorded.

Largely owing to imperfect preservation my attempts to obtain satisfactory
microscopic enlargements of the ornamentations of the interspacial areas of the
Estheriae:- dealt with in the present paper have not been successful; therefore I
decided not to make any special reference to these particular features of this
interesting genus so well represented in the Upper Permian rocks of the Newcastle
Coal measures of New South Wales.

ESTHERIA COGHLANI Cox. Plate ii, figs. 3, 4, 5.

Proc. Linn. Soc. N.S.W. 1880, p. 276 (1881); Etheridge, Junr., Mem. Geol. Surv.
N.S.W., 1888, Pal. No. 1, Pl. i, figs. 1-10.

To Mr. Etheridge’s description of this species nothing remains to be added.
He was of the opinion that among the drawings and specimens examined by him
there was a second species. On comparing his figures 1, 3, 6 with figures 8 and 9
that opinion seems to have good evidence to support it. After a careful study of
a large number of the specimens from the Hstheria beds, my view on the point in

question agrees with that of Mr. Etheridge, but available evidence is not yet quite
conclusive.

BY JOHN MITCHELL. 107

The range through geologic time of H. coyhlani was considerable, for at least
one specimen of it was obtained by me from the Wianamatta formation at Glenlee,
thus affording evidence of its persistence throughout the Triassic formations as
far as represented in New South Wales.

ESTHERIA IPSVICIENSIS, n. Sp. Plate iii, figs. 1-4.

Estheria mangaliensis Etheridae (nec Jones), Jack and Eth., Geol. and Pal. of
Queensland and N.G., 1892, p. 387.

The late R. Etheridge Junr. described this species but did not figure it. He
referred it to HE. mangaliensis Jones. His description of it is as follows: ‘Valves
broadly subovate, hardly convex; dorsal margin straight, about half the length
of the valves, terminated at the anterior end by the umbones; anterior, posterior
and ventral margins fully and broadly rounded, the anterior shorter than the
posterior, rendering the valves narrower at the former of the two ends; umhbones
depressed; ridges twelve in number, but probably two or three more exist on each
umbo, strong and well marked; interspaces wide, depressed, or perhaps very
slightly concave, especially towards the ventral portion of the valves; reticulation
not preserved. Length about three-sixteenths of an inch.”

“Observations: This little fossil, the first Hstheria found in the secondary rocks
of Queensland, appertains to the group represented by such species as
E. mangaliensis, E. Forbesi, etc., and in fact is so very close to the former in its
general features, that I am constrained to consider it as identical, notwithstanding
the absence of the reticulated surface in our fossil. It is particularly like
Professor Jones’ Pl. 2, fig. 16.”

In the above description and observations there appear to be some inaccuracies
which require correction; and to enable me to make these corrections I have
through the courtesy of Mr. B. Dunstan, Chief Govt. Geologist, Dept. of Mines,
Brisbane, before me the specimens used by Mr. Etheridge when making his
description and determination of the Hstheria under review.

In the first place the valves of the carapaces of this Hstheria are said to be
hardly convex, instead of being strongly so, when not flattened by external
pressure. The interspaces are neither wide nor depressed, but narrow and convex,
and flat near the ventral margin. Owing to the gradual way in which the dorsal
margin merges into the posterior margin it is difficult to give the exact ratio which
the dorsal margin of the Ipswich Estheria bears to the total length of its valves:
but the following measurements taken with care will result in a fairly accurate
estimate of the ratio being arrived at:

1. Length of dorsal margin and of valve 4: 7 (No. 55 in Coll. Mitchell).

2. Length of dorsal margin and of valve 5:12 (Type, Geol. Mus., Brisbane).

3. Length of dorsal margin and of valve 4: 9 (No. 53 Geol. Mus., Brisbane).

It may be noted here that in the case of three specimens of H. mangaliensis
Jones (Mon. Fossil Estheriae, 1862, p. 78, pl. ii, figs. 16, 20 and 21), the ratios of
the lengths of the dorsal margins and lengths of valves are 3:5, 1:3 and 7:18
respectively, exhibiting much variation. These proportions between the dorsal
margins and the lengths of the respective valves do not support the identity
of the Ipswich Estheria with the one from Mangali.

The anterior and posterior ends of the Ipswich Estheria are protrusively
rounded, and in this respect differ from the Mangali species. The umbones are
depressed only when distorted by pressure: ridzes visible on mature Ipswich LOPES)
are twenty-two or more. 7

108 FOSSIL ESTHERIAE OF AUSTRALIA, i.

I consider this Ipswich Hstheria to be a new species, and dedicate it to the
locality or measures from which it was obtained.

It differs from H. mangaliensis (1) in having a more ovoid form, (2) a greater
number of growth ridges, (3) in the individual specimens being more uniform
in outline, (4) in having more protrusive anterior and posterior margins. From
E. forbesi it is easily distinguished.

The illustrations of specimens of the species which accompany the present
paper, so clearly show the important features of the species that the writer
thinks it unnecessary to give further detaiis except as to dimensions, which are
as follows.

Plate iii, fig. 3, is from a specimen dealt with by Etheridge, who also examined
the specimens shown on plate iii, fig. 4.

1. Length 5:0: width 3:0 mm. (Type, Geol. Mus., Brisbane).
2. Length 5:5: width 4:0 mm. (No. 27 Geol. Mus., Brisbane).
38. Length 6:0: width 5:0 mm. (No. 55 in Coll., Mitchell).

Loc.—Denmark Hill, Ipswich, Queensland.

Horizon.—Upper ‘Triassic (Jack and Etheridge, Dunstan, Walkom and
Tillyard).

ESTHERIA GLENLEENSIS, n. sp. Plate ii, fig. 6.

Spec. chars.: Carapace transversely oval, dorsal margin mildly sagged, the
other margins rounded, the anterior strongly so; beak prominent about five-
eighths anterior, concentric striae nine or ten in number distinct, widely spaced;
the whole carapace convex.

Dimensions: Length 4:0; width 3:0, mm., approx.

This species is easily distinguishable from #H. coghlani. In some resvects
it resembles H. mangaliensis Jones (Mon. Foss. Estheriae, 1862, pl. ii, fig. 16), but
differs from that species in (1) having many less growth ridges, (2) having a
more truncate posterior margin, (3) a more protrusively rounded anterior margin,
(4) being of smaller size, and in having wider spaces between the growth ridges;
also in having a less variable outline.

The horizons in which the Mangali and the Glenlee Hstheriae are found do not
differ much in point of geologic time, therefore their identity would not have been
surprising, if such had been proven.

Loc.—The species was obtained from a cutting on the Great Southern Railway
near Glenlee Homestead.

Horizon.—Wianamatta series.

The fossils found associated with this Hstheria consist of other Hstheriae,
plants belonging to the genera Taeniopteris, Macrotaeniopteris, Thinnfeldia,
Cladophlebis and Phyllotheca, all of which go to show that the Wianamatta
Estheriae dwelt in fresh water.

ESTHERIA WIANAMATYENSIS, n. Sp. Plate ii, figs. 7, 8.

Spec. chars.: Carapace small convex, transversely subelliptic; beak sub-
central; ridges six or seven only visible, interspaces relatively wide; anterior and
posterior margins protrusively rounded dorsal margin arcuate, ventral margin
mildly rounded.

Dimensions.—Length 3:0; width (depth) 2:0 mm.

BY JOHN MITCHELL. 109

This may represent an immature specimen, but even if that be so, its tumid,
almost centrally placed beak and transversely elliptic shape, separate it from
other Australian species, and also from H#. minuta Alberti.

Loc. and horizon.—As for E. glenleensis.

ESTHERIA NOVOCASTRENSIS, n. sp. Plate iii, figs. 5, 6.

Spec. chars.: Carapace, outline subquadrate, dorsal margin straight, anterior
and posterior margins truncate, ventral margin widely and gently rounded; con-
centric striae about twenty in number, fine, evenly spaced; test appears to have
been of delicate texture, and is much crinkled; beak anteriorly situated and
prominent.

Dimensions.—Length 8; width 6mm. Another L=9 and width 6 mm. (These
dimensions are greater than for any other fossil Hstheria from the Newcastle
Series. )

I know of no species with which this can be either compared or contrasted with
advantage, though in the number of striae H. forbesi Jones, and the present form
approximate.

Loc.—Merewether Beach, a short distance S.W. of the outflow of the New-
castle sewerage; between low and high tide levels.

Horizon.—A few feet below the Dirty Seam of coal of Newcastle Coal Measures,
associated with various species of Glossopteris, and with Phyllotheca australis.
Upper Permian.

ESTHERIA LENTICULARIS, n. sp. Plate iii, fig. 7.

Spec. chars.: Carapace lenticular, small, smooth; beak subanterior, incon-
spicuous; concentric striae very fine six or seven in number, interspaces increase
in width as they approach the ventral margin.

Dimensions.—Length 2; width 1:8 mm.

The smoothness of the carapace and its small size are the two leading features
of this Hstheria; the fineness of its concentric striae is also very noticeable.

The only Australian Hstheria, which may be compared and contrasted with it,
is H. wianamattensis. They are both of small size and have few concentric striae
on their valves, but the species here dealt with has a more prominent and
anteriorly situated beak than the other one has. The fineness of the striae and
smoothness of the carapace of the present species strongly contrast with the
similar parts of FE. wianamattensis. It is not improbable that the Wianamatta
species descended from the mere ancient one from the Newcastle coa! measures.

Loc. and horizon the same as for HE. novocastrensis.

ESTHERIA TRIGONELLARIS, Dn. Sp. Plate iv, fig. 6.

Spec. chars.: Outline subtriangular, convex, dorsal margin slightly sinuate,
short; the other margins well rounded; beak about two-thirds anterior, very
prominent, elevated; concentric folds distinct, fairly regularly spaced, from the
end of the dorsal margin, the upper part of the posterior margin is depressed along
its edge. The description above is made from a mould of a left valve.

Dimensions.—Length 7:0; width 5:0 mm.

Loc. and horizon the same as for H. novocastrensis.

ESTHERIA OBLIQUA, n. Sp. Plate iv, fig. 1.
Spec. chars.: Transversely suboblong, convex, posteriorly obliquely protrusive,
dorsal margin long, straight anterior margin short, sharply rounded and gently

110 FOSSIL ESTHERIAE OF AUSTRALIA, i.

sinuate near the beak, posterior margin truncate; beak anterior and prominent;
concentric folds, strongly developed, twelve visible; but probably had sixteen or
more; they are strongly geniculated where they cross the umbonal fold, and as
they near the ventral margin, they are reduced to striae and are closer together.
The ornamentation apparently is made up of small globular bodies.

Dimensions.—Length 7:0; width 5:0 mm.

The specimen described has the valyes in apposition, the right being under
the left as may be observed by an examination of figure which represents the

type.
Loc. and horizon the same as for H. novocastrensis.

ESTHERIA LATA, n. Sp. Plate iii, figs. 8, 9.

Spec. chars.: Obliquely subflabellate, convex. smooth near the umbonal area;
dorsal margin straight, half as long as the length of the valves; anterior margin
distinctly rounded, posterior margin rounded and obliquely sub-protrusive, ventral
margin widely rounded. Concentric striaé. Fourteen visible, fine and fairly
distinct, and regularly spaced; beak inconspicuous, subanterior.

Dimensions.—Length 7-0; width 6-0 mm.

This is a very distinct species. easily separated from all other Australian
forms discovered up to the present time; also it ranks among the largest of the
fossil Estheriae from Australian palaeozoic rocks. In some respects it resembles
some forms of EH. forbesi Jones (Mon. Foss. Hstheriae, Palaeontological Soc., 1862,
p. 109, Pl. iv, figs. 8-9) in a general way; but specifically it is a distinct type.

Loc. and horizon.—Merewether Beach, near the Newcastle Sewerage outlet;
just below the Dirty Coal Seam, Newcastle Coal Measures, Upper Permian.

ESTHERIA BELMONTENSIS, n. sp. Plate iv, fig. 5.

Spec. chars.: The carapace subquadrate, flattish, dorsal margin straight,
relatively long. Under the dorsal line there is a flat area which resembles a
hinge plate, and which is not usually observed in members of the Estheria group;
anterior subtruncate, and the posterior one widely and mildly rounded; ventral
margin gently rounded, beak anterior, and inconspicuous, concentric striae, nine-
teen or twenty in number, distinct and fine.

Dimensions.—Length 6:0; width 5-0 mm.

This species in some features bears a strong resemblance to EH. ipsviciensis,
which may be a descendant of the older Permian type.

Loc.—The chert (Tuff) quarries near Belmont, Parish of Kahibah, County
Northumberland, New South Wales, associated with several species of Glossopteris,
Leaia and Insects.

Horizon.—Upper Permian.

ESTHERIA GLABRA, nn. Sp. Plate iv, figs. 2, 3.

Spec. chars.: Left valve testless, transversely subelliptic; convex; test was
apparently thin and delicate; dorsal margin very gently curved, anterior and
posterior margins rounded, ventral margin mildly rounded; beak, three-fifths
anterior, prominent; concentric ridges, only three or four clearly visible, but
traces of others can be seen with the aid of a good lens.

Dimensions.—Length 4:0; width 2:0 mm.

Although a large part of the cast of the valve above described has a smooth
appearance, there is slight evidence that the concentric striae on the original test

BY JOHN MITCHELL. P 1h lal

were more numerous than are at present visible. Its much elongated subelliptic
shape separates it from other Australian Hstheriae. The sparsity of its visible
striae is another distinguishing feature. The type is a unique specimen.

Loc. and horizon the. same as for EH. belmontensis.

ESTHERIA LINGUIFORMIS, n. sp. Plate iv, fig. 4.

Spec. chars.: Carapace obliquely flabellate, and flattened along the borders
of the postero-ventral margins. Dorsal margin straight and long, anterior margin
short and rounded; posterior one wide and gently rounded; beak anterior, incon-
spicuous; concentric striae about eighteen in number, obliquely directed towards
the postero-ventral margins, fine and compacted near the umbo and towards the
postero-ventral margins, intermediate of these two areas they are distinct or ridged.

Dimensions.—Length 5:0; width 4:0 mm.

The above description of the type is made from a mould of a nearly perfect
right valve.

This species, like EH. belmontensis, bears some resemblance to H. ipsviciensis,
so much so, indeed, as to make the assumption that the former is directly ancestral
to the latter, not an unreasonable one.

The distinguishing features of this form are (1) its anterior beak, (2) long
straight dorsal margin, (3) narrow anterior margin, and (4) wide posterior
margin.

Loc. and horizon the same as for H. belmontensis.

EstHEeRIA (?) BELLAMBIENSIS, n. sp. Plate iv, figs. 7, 8.

Spec. chars.: carapace sub-oblong, very inequilateral. Dorsal margin very long
and straight, anterior and posterior margins gently rounded, or subtruncate;
ventral margin subparallel to the dorsal one; beak, anterior, inconspicuous; con-
centric striae arranged in two sets—a wide apart set, and between each pair of
these, finer ones occur, very numerous; the vaives are convex.

Dimensions.—Length 10:0 mm.; width 6:0 mm. approximately.

If this fossil is an Hstheria, it exceeds in size all other Australian species of
the genus. In the character of its growth lines it resembles some Unionella; but
of three specimens known, each has a length of approximately 10-0 mm., and
appears to be mature. This length is much less than any mature Unionezila
possesses. The straight hinge line of the present fossil clearly separates it from
molluscans of that group. In shape, size and surface ornamentation H. bellam-
biensis resembles the H. striata group. The tength of the carapace of the species
under discussion exceeds that of all other Mstheriae known to me from palaeozoic
rocks.

Locality—The north side of the railway line connecting the South Bulli
(Bellambi) Colliery with the staiths at Bellambi roadstead, associated with
various species of Glossopteris.

Horizon.—About 150 feet below the seam of coal worked in the South Bulli
Mine, and in stratigraphical position near to the Belmont Insect and Phyllopoda
beds. Upper Permian.

My hearty thanks are extended to Dr. C. Anderson, Director of the Australian
Museum, Sydney, for placing at my disposal for examination a number of recent
Hstheriae. To Mr. W. S. Dun, Palaeontologist, Department of Mines, Sydney; and
to Mr. Booker, of the same Department, I tender sincere thanks for valuable aid
extended to me while preparing the present paper. I am thankful to Mr. H. G.

112 FOSSIL ESTHERIAE OF AUSTRALIA, i.

Gooch, Photographer for the Department of Geology, University of Sydney, for
some excellent photos of specimens dealt with in this paper, and lastly I am
indebted to my friend T. H. Pincombe, B.A., of New Lambton, for the privileges of
examining the Estheriae collected by Mrs. Pincombe and himself.

I wish to point out that in my paper on ‘Descriptions of New Species of Leaia’’
(These PROCEEDINGS, 1925, pp. 4388-447) I have made a mistake in orienting the valves of
the carapaces. I find that all right valves are designated left and the left valves right.

EXPLANATION OF PLATES II-IV.
Plate ii.

1. Deltopecten rienitsi, n. sp. right valve.

2. Deltopecten rienitsi, n. sp. left valve.

3. A right valve of Estheria coghlaini Cox somewhat pressed into the left valve. From
Cremorne bore. x 8. Coll. Dept. of Mines, Sydney.

4. Estheria coghlani from Dent’s Creek bore. x 10. Coll. Mitchell.

5. Estheria coghlani from Glenlee (Wianamatta Series). x 14. Coll. Mitchell.

6. Estheria glenleensis, n. sp. x 12. Coll. Mitchell.

7. Estheria wianamattensis, n. sp. xX 12.

8. Estheria wianamaitensis, n. sp. x 14.

Plate iii,
1. Estheria ipsviciensis, n. sp. Left valve. x 8. Coll. Mitchell. Presented by the Dept.
of Mines, Brisbane. 2
Estheria ipsviciensis, n. sp. Left valve. x 10. A good specimen. Coll. Dept. of
Mines, Brisbane, No. 53.
3. Estheria ipsviciensis, n. sp. Left valve (depressed). x i0. This valve was one
dealt with by R. Etheridge Junr. as H. mangaliensis. Coll. Mines Dept., Brisbane.
4. Estheria ipsviciensis. The photograph shows two individuals on the same stone.
They were examined by R. Etheridge Junr. (loc. cit.) x 10. Coll. Dept. of Mines,
Brisbane. :

bo

5. Estheria novocastrensis, n. sp. Left valve. x 5. Coll. Mitchell.
6. Estheria novocastrensis, n. sp. Left valve. x 5. Coll. Mitchell.
7. Estheria lenticularis, n. sp. x 12. Coll. Mitchell.
8. Estheria lata, n. sp. x 6. Coll. Mitchell.
9. Estheria lata, n. sp. Left valve. x 5. Coll. Mitchell.
Plate iv.
1. Estheria obliqua, n. sp. Left valve. x 10. Coll. Mitcheil.
2. Estheria glabra, n. sp. Left valve. x 12. Coll. Mitchell.
3. Estheria glabra, n. sp. A mould of right valve showing a more ovoid form than
Dives. 45 3 He
4. Estheria linguiformis, n. sp. Right valve. Coll. Mitchell.
5. Hstheria belmontensis, n. sp. x 7. Coll. Mitchell.
6. Estheria trigonellaris, n. sp. Left valve. x 7. Photo from a mould. Coll. Mitchell.
7. Estheria (7?) bellambiensis, n. sp. x 6.
8. Estheria (7?) bellambiensis, n. sp. A pair of fragmentary valves joined along the

hinge. x 6:5. Coll. Mitchell.
(The photographs by H. G. Gooch, J. A. Booker and J. Mitchell.)

A NOTE ON A DICOTYLEDONOUS FOSSIL WOOD FROM
ULLADULLA, NEW SOUTH WALHS.

By C. BarnarpD, B.Sc., Demonstrator in Botany, University of Sydney.
(From the Botanical Laboratories, University of Sydney.)

(Plates v and vi, and six Text-figures.)
{Read 30th March, 1927.]

Introduction.

The subject of this communication is a piece of fossil wood of a dicctyledon. It
was found by Miss Brown of the Geology Department, University of Sydney, in
a silica quarry at Bannister Head, near Ulladulla, New South Wales.

The author published a brief descriptive note of this fossil in connection with
Miss Brown’s paper on the Tertiary Formations on the South Coast of New South
Wales. It seems desirable that a fuller account of this interesting specimen,
supplemented by figures and photographs, should be recorded in order that its
systematic position may be ascertained.

Two fossil Angiosperm stems, with structure well preserved, have previously
been described from Australia. Sahni“? has published a description of two stems
from the Tertiary of Queensland. One, Petaloxylon scalariforme, from Mt.
Meershaum, near Nerang, is peculiar in that the chief conducting elements are
large scalariform tracheids. The medullary rays are 1-3 seriate, twenty-five to
thirty cells in height and pursue a sinuate radial course. The other, Petaloxylon
porosum, has essentially the same structure except that true vessels are also
present. Nobes™ records the discovery of four petrified stems from some Tertiary
Brown Coal deposits in South Australia. The preservation of these was so poor
that no description was published.

A great number of leaf impressions have been described by Httinghausen®
and Deane.” “” These remains, which have mostly been derived from Tertiary

1The silica beds, in which the specimen was found, were formerly believed to be of
Permo-Carboniferous age and the silicification was thought to have been effected by
the intrusion of volcanic dykes (Harper). ® 4° Miss Brown®) has lately proved that
these strata are of Tertiary age and has shown on reliable geological evidence that
Silicification has been effected by later Tertiary basalts.

These “silica rocks’ occur in patches, associated with clays and sandstones, in the
Milton-Ulladulla district on the South Coast of New South Wales, overlying beds of
Upper Marine age. At Bannister Head there is a slight angular unconformity between
the later sediments and the underlying Permo-Carboniferous strata; and also at
Pattimore’s Lagoon deposit, as well as at Bannister Head, remnants of the basalt flows
are evident in the form of capping sheets.

Petrographic studies by Miss Brown of the olivine basalt composing these lavas have
established the correlation between the flows in this region and other Tertiary basalts
of the State. Though the exact age of these ‘“‘Tertiary Olivine Basalts’” has not been
definitely determined, it is generally held (Sussmilch)@” that they belong to the Upper
Miocene or Pliocene period. The deposition of the underlying silica beds must then be
assigned to the Lower Tertiary Period.

O

114 DICOTYLEDONOUS FOSSIL WOOD FROM ULLADULLA.

deep leads, come from the same period as the Ulladulla fossil and give an indica-
tion of the Angiosperm flora existing at that time. .

Ettinghausen has described fruits as well as leaf impressions of Casuarina,
Persoonia, Grevillea, Hakea, Lomatia, Dryandra, Callicoma, Ceratopetalum,
Pomaderris, Eucalyptus, Santalum, Myrica, Quercus, Alnus, Acer, Aralia, Cinna-
momum, Fagus and Hlaeocarpus from Vegetable Creek, near Hmmaville, New
South Wales. These, it will be observed, include many endemic genera as well
as representatives of the Malayan flora.

The same author has described many types from Dalton, near Gunning, New
South Wales. Deane® ™ has identified a number of leaf impressions from the
clay beds at Mornington, Berwick, and Wonwron, Victoria; R. M. Johnston® “
describes similar impressions from the Lower Tertiary of Tasmania.

These remains have, with few exceptions, all been referred to existing genera.
In some cases the resemblance between fossil forms and living species is so close
that the affinity has been expressed in the specific name given to the fossil.
Hucalyptus pre-coriacea from the Lower Tertiary, for example, is almost identical
with the existing species “‘coriacea.’”’ From this evidence it will be observed that
the flora of the Lower Tertiary was very similar to that existing at present.

Text-figure 1. Representation of portion of the transverse
section (slightly diagrammatic), showing the seriation of the
vessels and fibres; illustrating particularly the thickness of the
walls of the vessels and fibres. x 280.

BY C. BARNARD. 115

Description.

Material. The specimen consists of a fragment of secondary wood preserved
by the agency of silica. It measures roughly 8 cms. by 4 cms. The end, which is
cut transversely, shows regular and well developed annual rings. The curvature
of these indicate that the piece of wood was about two and a half inches from
the centre of the stem and that the stem had a minimum diameter of five inches.
The structure of the wood is well preserved. Unfortunately the specimen does
not show any phloem or bark.

Microscopic Description.

Diagnosis: The wood has a fine texture. Annual rings developed. Vessels
numerous, circular, in radial series or isolated; bordered or scalariform pitting,
mostly porous perforate, tertiary bands present, slightly stratified, average
diameter 0-1 to 0-075 mm.; average of 50 per sq). mm. Wood fibres form the ground
tissue of xylem, thick walled, size variable, arranged in regular radial series.
Wood parenchyma cells abundant, rectangular, vertical height, 0:04 mm., width
0:25, occurring in vertical series. Medullary rays numerous, diffuse, biseriate,
rarely uniseriate, 10-15 cells in height, heterogeneous, cell walls thick with simple
oval or circular pits.

General: The vessels are of medium size and are evenly distributed throughout
the wood. The wood fibres compose the bulk of the wood and are very thick
walled. The vessels in the spring have a diameter up to 0:16 mm. These become
gradually smaller till at the end of the annual ring in the autumn the average
diameter is 0:04 mm. Parenchyma cells are abundant, diffuse and mostly situated
at the end of the annual rings, almost forming metatracheal bands. The medullary
Tays are very uniform in height. The vessels and wood fibres are arranged in
regular radial series.

Details of Elements:

In transverse section the Vessels show an approximately circular configuration.
They occur singly or in pairs or in radial rows up to ten in number. In these
latter cases the tangential walls are flattened (Text-fig. 1 and at 0 in Fig. 1,
Plate v). The average diameter is 6-075 mm. The average diameter of the spring
elements is 0-13 mm. and the autumn average is 0:05 mm. The walls are thick.
In longitudinal section the sculpturing on the walls consists of regular series of
small oval bordered pits in horizontal rows (Text-fig. 4c and Fig. 2, Plate v).
The borders of the pits are narrow.

The large vessel in the centre of Fig. 2 Plate v shows this type of pitting
very well. In some cases the coalescing of the pits gives rise to a close scalariform
sculpture. This feature is apparent in the large vessels in Fig. 3 Plate vi and
appears, in fact, to be the dominant type. The internal spiral bands of thickening,
termed tertiary thickenings, are also present in many of the vesséis.

The average length of the vessel segments is 0:6 mm. The two segments,
illustrated in Fig. 3 Plate vi, are typical and each of these has a length of 0-6 mm.
The majority are of the porous end-wall type (Text-figs. 4c and 6 and Fig. 2
Plate v and Fig. 3 Plate vi). The septa in these elements are short and distinct;
they are almost at right angles to the longitudinal walls of the vessel. Each is
perforated by one large pore. A few vessels exhibit a number of elongated pores
on the end-wall. These are examples of the scalariform perforate type. In some

116 DICOTYLEDONOUS FOSSIL WOOD FROM ULLADULLA.

eases the so-called “imperfect perforation’ is present. The end-walls in vessels,
which show this feature, are very oblique and numbers of bordered pits occur on
them together with one or two large perforations (Text-fig. 4 a@ and db). Vessels .
with end-walls of this nature are longer and have a smaller diameter than the
neighbouring vessels with the porous type. No tyloses were observed. The
vessels appear to be more or less stratified or storied.

2

Text-figure 2.—Representation of portion of the radial
longitudinal section showing the vessels in detail. An indication
of a close scalariform sculpturing is apparent on the lateral
walls of several porous end-wall types of vessels. x 80.

Text-figure 3 represents a portion of the tangential
longitudinal section showing the medullary rays in detail
together with vessels and fibres. On the left there is a series
of parenchyma cells with the original thickness of the walls
preserved. x 165.

The wood fibres vary considerably in diameter and length. The average
diameter is 0:02 mm.-0-01 mm. Very often the tangential diameter is greater than
the radial. In general the walls seem thin (Text-fig. 2), but this appearance is due
to the fact that it is really the junction line between contiguous fibres which
represents the wall. In places where the preservation of the structure iS more
perfect and the true thickness of the wall is shown it presents a very characteristic
appearance. The nature of the fibres is seen at a in Fig. 1 Plate v. The lumen

BY C. BARNARD. 117

represents only one-third or one-fourth of the diameter. No pitting could be
observed in the longitudinal aspects of these elements. They are regularly seriated
in a radial direction.

The wood parenchyma cells are seen to best advantage in the longitudinal
sections (Text-figs. 3, 4 and 5 and Fig. 3 Plate vi). They occur in vertical series
and are most abundant in the region of the autumn wood. There are as many
as forty cells in ome series. The average vertical height of each cell is 0:040 mm.
and the average width in tangential and radial aspect is 0-025 mm. The pitting
on these cells was not observed. The walls are comparatively thick. In some
instances black carbonaceous material is apparent in the cell cavity.

A

Text-figure 4.—Illustrating the types of end-wall found in
the vessels in radial longitudinal section. In a and 6 the end
wall is of the imperfect perforate type. At c is shown a typical
porous perforation in a vessel, which shows multiseriate pitting
on the vertical wall. x 280.

The medullary rays are numerous and diffuse, heterogeneous and without
exception uniseriate or biseriate. The vertical height of the ray is also a definitely
characteristic feature. They are relatively short. The biseriate type of ray is
predominant and in tangential section generally contains 20 to 30 cells (Text-figs.
3 and 5 and Fig. 4 Plate vi). These rays are therefore about 10 and 15 cells in
height. The two rows of cells, comprising the width of the ray in tangential
view, alternate. The uniformity of the width of the ray is one of the most
interesting features of the stem. The individual cells have a vertical height of
0:02 mm. and are approximately 0:07 mm. in radial length (Text-fig. 6). In

118 DICOTYLEDONOUS FOSSIL WOOD FROM ULLADULLA.

tangential view they appear rounded with an average diameter of 0:02 mm. A
large amount of black carbonaceous matter is present in the ceil cavity (Fig. 4
Plate vi). In places the pits on the walls of these cells may be distinguished in the
radial longitudinal section. They are more or less rounded and simple. The walls
of the ray cells contiguous to other ray cells are thick and are pitted in the same

manner. These pits could be observed in section as well as in surface view
(Text-fig. 6).

Text-figure 5.—Representation of portion of the tangential
longitudinal section of the autumn wood showing an abundance
of wood parenchyma series. x 165.

Systematic Position.

Our knowledge, at present, of the systematic anatomy of the dicotyledonous
stem is meagre. On this account the affinities of a fossil stem may be satisfactorily
determined, only when very distinctive features are observed.

Fliche,” Lignier,“ Stopes and Fujii,“® Stopes,°® Krausel,“® Bailey,™ and
Edwards, have described dicotyledonous stems from the Cretaceous and Tertiary
in other parts of the world. Hach of these authors has named the fossils described,
after the woods of the living genera, which they most closely resemble. In this
way the affinities of the specimens have been indicated. In the case of the
Ulladulla fossil it has been found that the structure indicates an affinity with a
certain Natural Order and not with one particular genus. There is a very close
resemblance between the woods of the various genera of this Natural Order. On
this account it is not possible to select one genus, which has a preeminent affinity
with the fossil. The writer therefore has not followed the precedent established
by the authors quoted above, but deems it wisest only to indicate the apparent
systematic position of the specimen.

The characters of every Order upon which information is available“ have
been carefully considered. Representatives of certain orders show similarities to
the fossil but differ in important details. Types of the Schizandreae group of the
Magnoliaceae, for example, possess a very similar combination of wood structures,

BY C. BARNARD. 119

but differ in the fact that the medullary rays characteristically have a great
vertical height. Certain genera of the Natural Orders Bixineae, Euphorbiaceae,
Anarcardiaceae, and Rutaceae have somewhat similarly constituted woods. These
particular genera, however, are not represented in the present-day Australian flora.
As a most striking resemblance is evident between the wood of the fossil and
the woods of an Order, which is widely distributed in Australia, it seems that
the similarities referred to above may be disregarded.

The writer has examined figures, descriptions or actual sections of repre-
sentative woods of most of the natural orders found in Australia. The
Saxifragaceae alone possess the same type of wood structure as that exhibited by
the fossil. The combination of wood structures, which are characteristic of this

Text-figure 6.—Diagrammatic representation of a radial
longitudinal section. All the features depicted may be clearly
observed in certain places in the sections. The heterogeneous
nature of the medullary ray is shown and the nature of the
pitting on the various elements. x 280 approx.

order, seems to be identical with the combination of structures present in the
fossil. The regular radial seriation of the vessels, the occurrence of scalariform
perforate end-walls in the vessels, as well as the narrow, heterogeneous, and
uniform medullary rays are characteristic features of the woody genera of this
order. For these reasons the writer tentatively places the Ulladulla specimen in
the Natural Order Saxifragaceae.

I am indebted to Mr. Welch, of the Technological Museum, for allowing me to
examine photo-micrographs of the woods of certain representatives of this order,

120 DICOTYLEDONOUS FOSSIL WOOD FROM ULLADULLA.

which I could not easily obtain in the living state. Weinmannia, Ackmana,
Schizomeria, Quintinnia, Callicoma, and Ceratopetalum show a remarkable
resemblance to the fossil. In the case of Ackmana the structures were almost
identical.

In conelusion, I wish to thank Professor A. A. Lawson for advice in the
preparation of the text, and Mr. Welch for the above-mentioned assistance. I am
indebted also to Dr. Carl Boesen and Mr. R. Murray, B.Sc., for the photo-
micrographs reproduced.

Summary.

1. The specimen described was obtained from the “Silica” beds at Bannister
Head, near Ulladulla, on the South Coast of New South Wales. These beds are
of Lower Tertiary age.

2. The specimen is a piece of silicified secondary wood.

3. The structure of the wood is that of a typical dicotyledon, and shows a
very close agreement with that of the Natural Order Saxifragaceae. The author
tentatively places the specimen in this Order.

I have endeavoured to represent the nature of the individual elements
accurately though a certain latitude has been taken in making the drawings.
In Text-fig. 2, for instance, the medullary rays have been omitted in this radial
longitudinal representation in order that the vessels may be seen more clearly.
Further a uniform state of preservation is not present throughout the specimen;
and irregularities in preservation render it impossible to depict the true character
of all the elements in a drawing with the camera lucida. The less obvious details
of structure in the various elements may not all be observed in one place in the
section. For these reasons it was thought advisable to draw together features
which could be definitely recognized into two diagrammatic figures (Text-figs.
1 and 6).

References.

® Barttey, I. W., 1924.—The Problem of Identifying the Wood of Cretaceous and later
Dicotyledons. Annals of Botany, Vol. 38.

® Brown, IpA, 1925.—Some Tertiary Formations on the South Coast of New South
Wales—with special reference to the age and origin of the so-called Silica rocks.
Proc. Roy. Soc. N.S.W., Vol. LIX.

@ DEANE, H., 1902.—Geol. Survey Victoria, Vol. I, Part I.

(4) —_______. 1902.—-Notes on Fossil Leaves from the Tertiary Deposits of Wingello and
Bangonia. Records of the Geol. Survey, N.S.W., Vol. VII.

©) ETTINGHAUSEN, C. VON, 1888.—Contributions to the Tertiary Flora of Australia.
Memoirs Geol. Surv. N.S.W. Pal, Series No. 2.

® Epwarps, W. N., 1925.—Report on Fossil Wood from Somaliland. Pal. Section,
Botanical Abstracts, 1925.

@ FLIcHE, P., 1905.—Note sur des bois fossiles de Madagascar. Bull. Soc. Geol. France,
Ser. 4, Vol. 5.

() Harper, L. F., 1916.—Annual Report, Dept. of Mines, Sydney.

(9) ________, 1918.—Annual Report, Dept. of Mines, Sydney.

ao) ________. 1924,__Dept. of Mines, Sydney, Geol. Survey. Bull. No. 10. Silica.

GD JOHNSTON, R. M., 1885.—Proc. Roy. Soc. Tasmania.

G2) ________, 1 §89.—-Proc. Roy. Soc. Tasmania.

G3 KRAUSEL, K., 1924.—Reference in the Palaeobotanical Section of the Botanical
Abstracts, 1924.

“4 LIGNIER, 1907.—Vegetaux fossiles de Normandie, Bois dive. Mem. Soc. Linn.
Normandie, Vol. 22.

45) Nopes, DorotHy, 1922.—A Preliminary Note on Fossil Woods from some Australian
Brown Coal Deposits. Vrans. and Proc. Roy. Soc. S.A., 46.

BY C. BARNARD. PAL

a6) SAHNI, B., 1920.—Petrified Plant Remains from the Queensland Mesozoic and
Tertiary. Queensland Geol. Survey Publication, 267.

G7) SOLEREDER, H., 1908.—Systematic Anatomy of the Dicotyledons. Trans. Boodle and

Fritsch, 2 Vols.

a8) Stopes, MARIE C., and Fusir, K., 1909.—Study on the Structures and Affinities of
Cretaceous Plants. Phil. Trans. Roy. Soc. London, Series B, Vol. 201.

ao) Stopes, Marig C., 1915.—The Cretaceous Flora. Part 2, Lower Greensand Plants of
Great Britain. Cat. of Plants of the Mesozoic in the Brit. Museum.

@o SUSSMILCH, C. A., 1923.—Presidential Address, Proc. Roy. Soc., N.S.W., Vol. LVIII.

EXPLANATION OF PLATES Vand VI.
Plate v.

1. Photo-micrographic representation of portion of the transverse section. The vessels
are clearly seen and in places the wood fibres are visible. Fibres are well shown
at a. The cellular structure of the numerous and narrow medullary rays is not
distinguishable. These cells are filled with carbonaceous material. x 200.

Photo-micrographic representation of radial longitudinal section. The principal
feature is a large vessel showing a large terminal porous perforation and multi-
seriate pitting on the lateral wall. The wood fibres may be seen and also
indications of the medullary rays, which are poorly preserved in this portion of
the stem. x 200.

bo

Plate vi.
Photo-micrographic representation of a radial longitudinal aspect showing two large
vessels with porous perforations. Medullary rays are seen crossing the vessels.
On the lateral walls of the vessels portions of the scalariform type of sculpturing
are discernible. x 200.
4. Photo-micrographic representation of the tangential longitudinal section. This shows
the medullary rays. At aa series of wood parenchyma is evident. x 200.

(J)

P

ON A CASE OF NATURAL HYBRIDISM IN THE GENUS GREVILLEA
[N.O. PROTEACEAE].

By C. T. Musson and the late J. J. FLrercHer, M.A., B.Sc.?

(Plate vii.)
[Read 30th March, 1927.]

In the Abstract of Proceedings dated 30th July, 1913, is given a short account
of a case of hybridism between two species of Blue Mountain Grevillea by the
late Mr. J. J. Fletcher. He contended, with many facts, much detail and a large
range of specimens, that G. Gaudichaudii, R.Br., consists of a series of transitional
forms between G. laurifolia, Sieb., and G. acanthifolia, A.C. That the entire series
known as G. Gaudichaudii is a series of variable naturally related forms, explain-
able only as being hybrids between the two other species mentioned. That the
two parent-species are markedly contrasted in most of their morphological
characters, in their habit: of growth, in being members of two different plant
associations, consequently in their habitats; cross-pollination being possible, how-
ever, by reason of comparative proximity, and because the racemes of both are of
the same pattern (elongated, secund and centripetal); moreover, the conditions
favouring cross-pollination arise only at or close to the boundary between their
respective habitats; consequently they are usually found in company with one
or other, or both the parent-forms.

G. Gaudichaudii has no specific characters; being of mixed origin, the
characters are inherited, being blends or mixtures of those of the parent-forms.
They are the first Australian, Proteaceous. wild hybrids recorded. They are not
infertile, but they rarely reproduce themselves, because likely to be overlooked by
visiting birds (the natural agents in pollination) or are likely to be pollinated
from one of the parent-forms. (C.T.M. and J.J.F.)

Historical.

During the stay of the French corvette Uranie, engaged in a scientific voyage
round the world, in command of Captain Freycinet, in Port Jackson, in November
and December, 1820, Gaudichaud, the botanist of the expedition, made a fruitful
excursion to Bathurst. Among the most important botanical additions to his
collection was a rare, undescribed Grevillea, probably the only plant of its kind
seen by him. It was obtained on the Blue Mountains, at “Vallée de Jamison.”

‘This paper is the outcome of an examination of extensive notes on the subject left
by the late J. J. Fletcher and has been prepared for publication at the request of Mrs.
Fletcher. Having had the opportunity to assist in the field work, knowing Mr. Fletcher’s
views, having examined the species in the field and worked over the collected specimens,
I can endorse fully the detail and conclusions given. Except where indicated, this paper
is copied directly from Mr. Fletcher’s notes.—C. T. Musson.

BY C. I. MUSSON AND J. J. FLETCHER. 123

But Gaudichaud did not go far enough from the Western Road to see the Jamison
Valley which, at that time and for long afterwards, was inaccessible from the
main road. What he means, we may fairly well conclude, was the little valley and
its tiny rill of water visited by C. Darwin.* (It seems quite probable that
Gaudichaud when at The Weatherboard on his way to Bathurst teok advantage
of the short time at his disposal to take the (probably) one walk that presented
itself, and which was later taken by C. Darwin. This is an interesting conjecture,
and is quite likely correct, for in those early days there were no side roads opened
up and any visitor would doubtless be directed, when the coach stopped at The
Weatherboard, the old name for Wentworth Falls, to the one track, apparently
between what is now known as King’s Tableland, and the more westerly spur at
the end of which now exists the Wentworth Falls Hotel. The tiny creek runs
approximately south from the township and falls over the cliff as described by
Darwin. Gaudichaud could not have collected his plant in the Jamison Valley;
but on the high ground overlooking it; somewhere along the valley mentioned
above.—C.T.M.)

On his return to Paris, Gaudichaud gracefully presented a portion of his
specimen to Robert Brown, who subsequently described it in the Suppl. Prod. (1830)
under the name of G. Gaudichaudii, R.Br.

In the meantime the scientific records of the voyage of the Uranie had been in
preparation and Gaudichaud wished to include 2 description of his new Grevillea
in the botanical part of the work, which was eventually published in 1826. This
description, the earliest to be published, is usually quoted as if it were R. Brown’s.

From a comparison, internal evidence shows it is evidently Gaudichaud’s own
description that he gives. Brown may or may not have sent him a copy of his
proposed description. But as Gaudichaud’s specimen was not exactly a duplicate
of the one he had parted with, and he wished to figure it, he evidently described
his own specimen, adopting R. Brown’s name.

Forty years later Bentham redescribed G. Gaudichaudii more fully than had
his predecessors (Fl. Aus., V, p. 488); but unfortunately for those who came after
him, he apparently did not see R. Brown’s specimen, nor specimens like them and
in drawing up his description he omitted to make provision for any but his own.
The consequence is that both the status and the identity of G@. Gaudichaudii are
left in doubt. If we are to take Bentham’s description as the standard for deter-
mining G. Gaudichaudii, then the specimens of R. Brown are not that species,
and we are given no clue as to how they are to be disposed of. It seems remark-
able that Bentham apparently did not himself realize this anomaly.

Modern Details.

At the meeting of this Society in July, 1910, in response to an invitation issued
at a previous meeting asking members who had collected, or who had in their
possession specimens of G. Gaudichaudii, four members brought specimens, some

2“Tn the middle of the day (1844) we baited our horses at a little inn called
The Weatherboard. The country here is elevated 2,800 feet above the sea. About a
mile and a half from this place there is a view exceedingly well worth visiting. Following
down a little valley and its tiny rill of water, an immense gulf unexpectedly opens
through the trees which border the pathway, at a depth of perhaps 1,500 feet. Walking
on a few yards, one stands on the brink of a vast precipice, and below one sees a grand
bay or gulf. . . . About 5 miles distant in front, another line of cliffs extends, which
thus appears completely to encircle the valley.’-—Darwin’s Naturalist’s Voyage Round the
World, 1889, p. 523.

124 NATURAL HYBRIDISM IN GENUS GREVILLEA.

of them from the Botanic Gardens Collection. The joint collection then exhibited

was, almost certainly, the best at the time in Australia. Three collectors had
obtained plants which answered to the descriptions of Brown and Gaudichaud;
but no plant was produced which corresponded with Bentham’s description. Since
‘then I have devoted my brief holidays to the further study of these plants under
natural conditions, and have now (1915) a much better collection than that
exhibited in 1910; but I have not yet found a single plant to which Bentham’s
description will apply. Nevertheless I raise no doubt as to the correctness of
Bentham’s descriptive details, though I think one of his inferences was made under
a misapprehension, due to want of knowledge not determinable from the examina-
tion of herbarium specimens alone; and that this is open to correction. I believe

I am now in a position to offer a solution of the difficulties which arise from the
conflicting descriptions.

The Conflicting Descriptions.

The three descriptions mentioned, together with a note by Mr. J. H. Maiden,
which was not intended to be descriptive but to put on record some observations
of the late Mr. W. Forsyth! on living plants, comprise all the literature about
G. Gaudichaudii available.”

Brown refers to some MS. notes of Allan Cunningham’s, at the end of his
description in which the species is referred to under the MS. name G. acanthifolia
var. quercifoliae. Unfortunately Cunningham’s notes were not published. There is
no mention of the species in his “Specimen of the Indigenous Botany of the
Mountainous Country between Sydney and Bathurst” (Field, 1825), though
G. acanthifolia was first described in this paper; or in any other of his published
writings as far as I know.

Dr. Woolls and Baron von Mueller, as far as I can ascertain, never found any
occasion to refer to it.

In order, therefore, to make clear how some specimens I have now collected
come in, in an important way, I preface an account of these with a brief statement
of the whole case, based chiefly upon a comparison of the descriptions of
G. Gaudichaudii by Brown, Gaudichaud and Bentham, supplying necessary
comments.

Though not first in chronological order, I begin with R. Brown’s description.

One minor correction needs to be made, otherwise the description is satis-

.factory as far as it goes; having no knowledge of the habit of G. Gaudichaudii, he
naturally took the spikes to be erect. The plants are prostrate, as pointed out in
Mr. Maiden’s note, and the spikes are more or less horizontal as in G. laurifolia;
and, as in that species, frequently rest upon the ground.

From a comparison of the descriptions of G. Gaudichaudii and G. acanthifolia,
it is evident that R. Brown considered Gaudichaudii to be distinguishable from
acanthifolia by the scattered appressed hairs on the underside of the leaves, by
their simpler lobing, by the presence of an intra-marginal vein, by the racemes
not being so dense and by the perianths having less clothing (sericeous as
compared with very villous). The term sericeous, however, in the specimens I have
seen applies only to the white silky hairs; but besides these there is some
hairiness tinged with the colour, dark crimson, of the perianth as in G. laurifolia.
If the white silky hairs could be removed the perianth would not be glabrous.

1 These PROCEEDINGS, 1904, p. 749.
2 Written between 1913 and 1918.

BY C. T. MUSSON AND J. J. FLETCHER. 125

From my point of view, the condition in question is the result of a mixture
of the corresponding characters of G. lawrifolia and G. acanthifolia when these two
species are crossed.

One other point, the lobes of the pinnatifid leaves are described as entire, a
character in which R. Brown’s specimen differed from that described and figured
by Gaudichaud. Portions of plants showing this character may be procured
without difficulty, but in all plants of the Gauwdichaudii group (see ii below) that
I have seen, if all the leaves are examined, some of them will be found to offer
indication of secondary lobing. Brown’s and Gaudichaud’s specimens may, there-
fore, have been different portions of the same plant, and yet offer the differences
recorded. Finally R. Brown definitely recognized that the racemes and flowers
were not exactly like those of G. acanthifolia.

Turning to Gaudichaud’s description (he saw both species in the living
condition), it is evident he did not copy the descriptions of Brewn, but drew up
his descriptions independently. He recognized, like Brown, the differences in the
indumentum of the leaves, in the lobing of the leaves, in the clothing of the
perianth (sericeous in one case, tomentose in the other), but he used the wrong
adjective in both cases in describing the pistils; the styles are glabrous in both
cases, but the ovaries are villous in both cases.

A very interesting and important omission, however, is that of the presence
of the intra-marginal vein. If he had had the opportunity of seeing the leaves of
G. laurifolia (which he apparently did not collect), its presence might have struck
him. From this omission, and the mistake about the pistils, as well as in other
small details, it seems clear to me, that R. Brown did not write this description,
and that in all probability Gaudichaud did not see a copy of R. Brown’s before his
own was published. He gives a good figure of a portion of a stem or branch
with nine leaves, having 5, 7, 8 and 9 lobes; and two spikes which are evidently
not exactly like those of G. acanthifolia; whilst the ovary is correctly figured as
villous (Frey. Voy. Bot., p. 443, and Plate 46).

Turning now to Bentham’s description it appears that the plants to which
it applies are not known to local collectors; it may be that they are a second
cross between one of the ordinary forms and a hybrid.

Some Indirect Evidence.

In the neighbourhood of Sydney it is easy to find plants of two or more
different species of Grevillea flowering at the same time and fruiting freely,
growing close to each other under conditions apparently quite favourable to cross-
pollination by nectar-seeking birds. Nevertheless transitional forms between such
species have never been reported. They do not seem to be produced in the cases
examined, apparently for reasons given below.

For some years I have had an excellent opportunity of periodically inspecting
an extensive crop of thousands of specimens of the two species G. sericea and
G. buczifolia, the former more numerous than the latter, crowded together on a
ridge overlooking the Lane Cove River, 200 to 300 yards wide, it terminates
abruptly on the east. Half a mile west the Hawkesbury sandstone is overlaid by
Wianamatta shale, with a corresponding change in vegetation. This ridge was
swept by a destructive bush fire on 2nd January, 1909, a phenomenally oppressive
day with hot westerly wind. In due course, after rain had fallen, seedlings came
up in great profusion, especially so Grevilleas, which grew rapidly, got ahead of
the more slowly growing plants and temporarily took charge of the area. So

126 NATURAL HYBRIDISM IN GENUS GREVILLEA.

numerous were they that one could walk down one side of the ridge and up on
the other or zig-zag in any direction and yet keep one hand on one or more plants
all the time. The two species mentioned were flowering side by side, their branches
overlapping, so that the racemes of one were sometimes only a few inches from
the other. Seven capsules to a raceme were quite a common occurrence. The
plants are so different that they are readily distinguishable at sight at any stage
of growth. I was interested in the renascence of the vegetation on this circum-
seribed spot, having had the Grevilleas continuously under observation from the
time they were seedlings, yet I never succeeded in finding a single plant that was
not certainly determinable as one or the other. There appeared no evidence
whatever of any kind of successful hybridization, nor was there, as far as I have
observed, in a fresh crop of seedlings that came up. I am therefore led to
conclude that the Honey-eaters cannot cross-pollinate them.

Elsewhere G. punicea and G. buaxifolia, G. sphacelata and G. sericea; and on
the lower part of the Blue Mountains G. nhylicoides and G. parviflora, may be
found growing together under conditions which seem to be favourable for crossing,
but apparently without any such result. It may be that the pairs of species
enumerated are infertile, inter se; that is a matter for experimental investigation.

There is another possible explanation, which may he the correct one. Reference
to the Flora Australiensis shows that the species of every pair referred to are
placed in different sections of Bentham’s Table; in other words the racemes are
of different patterns or there are structural peculiarities of some sort present in
one and not in the other. The result probably being, that a nectar-seeking bird
visiting the flowers of one species may carry the pollen away from flowers on one
part of the head, their position not matching with that of the receptive stigmatic
surface of the other species. The result would be that cross-pollination of other
flowers in the same or in other racemes of one type could take place, whilst in other
species with raceme types differing it could not be effected.

Experimental investigation into these interesting details is much to be
desired. At present we have little or no data for consideration.

(It has been observed that Honey-eaters commonly make a practice of com-
mencing with the lower flowers of a raceme, working upwards. This would favour
cross-pollination in the case of laurifolia and acanthifolia, as in both species the
flowers open centripetally. In the case of sericea and buzifolia, one opens
centrifugally, the other centripetally, a fact which would tend to prevent crossing.—
C.T.M.)

Some Detail as to the Three Species from Fresh Specimens, 1913.
G. laurifolia (Plate vii, fig. 1.)

G. laurifolia (Plate vii, fig. 1) belongs to the xerophytic plant association
which successfully occupies the poor soil of dry situations in the Hawkesbury
Sandstone series (approximating to the Bunter Sandstone of the Trias). It
certainly occupies some very dry positions on the ridges or on more level areas
on the top of the tableland, where the ground must be so sunbaked and hard in
droughty summers, that when heavy rain falls comparatively little can be stored
as it runs away so rapidly. On the other hand mountain mists, which are not
infrequent, may be helpful.

These plants are prostrate in habit, with several radiating procumbent stems
arising from a thickened base. These stems are often 8-10 teet long, wiry and
flexible, and readily made up into a small coil (except when very old); they are

BY C. T. MUSSON AND J. J. FLETCHER. sey

well branched and very leafy, forming carpets lying close to the ground. In very
old plants the basal thickening forms a large knob and the stems arising from it
may be an inch in diameter or more for some feet from the base.

The leaves haye a characteristic venation and indumentum, and cannot be
mistaken for any other local plant. They are distinctly thicker than those of
G. acanthifolia. Petiolate, entire, they vary a good deal in shape and size,
according as they are well exposed to the light, or shaded by grass, shrubs or trees.
They may be ob-lanceolate, long and narrow, short and broad or broadly ovate,
even almost circular, and mucronate. Glabrous above when adult, very young
leaves and shoots are thickly coated with a mixture of silky white and ferruginous
or reddish appressed hairs lying very close together. Those of the upper leaf
surfaces are soon lost. Except the midrib the under surface is thickly coated
with silky white hairs lying close together and doubtless of use in checking trans-
piration as well as serving to protect the stomata from dust, or soil washed down
by rain. The veins are very conspicuous, nearly parallel on each side of the
midrib. They end distally in a characteristic well-marked intra-marginal vein.

The perianth tube, viewed from outside, has much more colour than in
G. acanthifolia, and this is all the more evident because most of the appressed
hairs clothing the tube are dark crimson or ferruginous, only a few white hairs
being mixed with them. The hairs appear as if plastered down, not merely
appressed. Racemes of very young unopened buds, with the bracts in situ, may
have the globular limb, and that part of the limb not hidden by the bracts,
ferruginous or rusty; or they may be turning crimson, though with a distinctly
ferruginous tint on the globular limbs. The bracts are yellowish, coated with
yellowish or rusty and white hairs. The rhachis is also thickly coated with short
crimson, ferruginous or rusty appressed hairs. As the tube dries the colour some-
times becomes almost magenta. In still older buds in which the bowed style has
not yet begun to protrude from the flowers, the tubes of the perianths, and the
pedicels, are more distinctly crimson, but the globular limbs are still ferruginous,
to the naked eye contrasting in colour. The rhachis may have more evident
crimson hairs mixed with ferruginous or rusty hairs; while the bracts may be
yellowish, or tinged with crimson near the base or over the greater part of their
surface. In still older racemes, in which the bowed styles of the lowest flowers
are protruding, there is still a noticeable contrast between the colour of the
tubes, and of the globular limbs; and so much of the inner surface of the limb
as is exposed, is seen to be edged with dark crimson, in still older flowers
changing to purple. This may be confined to the free edge of the split perianth
leaving a yellow gutter between, or the whole of the exposed inner surface may be
purple, becoming darker as the flowers mature; eventually fading to some shade
of dark red in dried flowers.

In mature flowers the globular limb and tube still show the difference—
crimson—ferruginous. The pedicels are same colour as the perianth—rusty. In
some racemes the flowers remain ferruginous without change, in others one-half
longitudinally is crimson, the other ferruginous. Stipes of the ovary is longer than
in G. acanthifolia. Styles lighter crimson, stigmatic disc green, sometimes crimson.
Freshly exposed pollen is bright yellow and floury, contrasting strongly with the
crimson. Hairs on the-ovary and the stipites not so numerous as in G. acanthifolia,
usually white mixed with crimson, or they may be nearly all crimson with just a
few white. I think the hairs of the tomentum are correctly described as appressed
all over (on tube and limb). The perianth and globular limb split on the lower

128 NATURAL HYBRIDISM IN GENUS GKEVILLEA.

side (Bentham), the latter nearly to the base to allow the exit of the pollen-
carrying disc, while the two laminae of the limb usually cohere on the upper
side. When exposed in this way the inner surface is purple or there is a purple
stripe on each side with an intervening longitudinal yellowish area along the
coherent margin extending for some distance below the revolute limb but not
to the base. The purple colour is evanescent, changing to dark crimson.

G. acanthifolia. (Plate vii, fig. 6.) é

This species is tolerant to any amount of water applied to the roots; it
belongs to a mesophytic assemblage of plants which flourish in wet swampy areas
mainly caused by soakage from springs. Such areas are noticeable from a distance
because of the absence of trees, and their green appearance. ‘They are often
mentioned by early explorers as they offered chances of feed for their horses
or bullocks when there was nothing to be had elsewhere. They occur at different
levels, frequently on the slopes of shallow valleys or bordering creeks,’ sometimes
they surround small islands of drier ground on which a few trees may occur.
They are to be found only in the upper portions of the tributary valleys of the
Grose and Cox rivers. Further down the valleys are enclosed between precipitous
cliffs, and this, with absence of sun, profoundly alters the conditions for plant life.

In very dry seasons these areas dry up on the surface and are often burnt
over. If the Grevilleas are only well scorched, the portions above ground die,
but after sufficient rain they again send up shoots from the thickened base.

G. acanthifolia also flourishes on the banks of creeks, close to the water; and
it is not unusual to see some of the main roots actually trailing in running water.
It occurs also in depressions near and along the course of creeks, supplied from
the overflow after heavy rains.

It sometimes occurs in unusual situations. I have seen one flourishing on
the side of an apparently dry embankment leading to a bridge crossing a creek,
and growing close to a plant of G. laurifolia. Possibly when rain falls the embank-
ment is capable of absorbing more moisture than one might expect; perhaps much
more than the sun-baked virgin ground which had never been stirred by spade or
plough, but in which Jauwrifolia can flourish.

Notes on Fresh Specimens, March and November, 1918.

Leaves: Trifid leaves sometimes subtend an inflorescence. Have seen 3-19
lobes; intermediate numbers commonest. There is an intra-marginal vein (or
perhaps an epidermal thickening) to the lobes, but it is not visible on the under-
surface; nevertheless, when held up to the light, it is visible like the ordinary
veins (anatomical examination is wanted here). Seedling leaves, and leaves of
side shoots, also the lowest leaves near the ground, of adult plants have relatively
much more lamina, and are more easily pressed flat, not being so rigid.

Shoots: Young shoots, and unexpanded young leaf-masses, are coated with
white silky hairs on both sides but not the upper surface of the young leaves,
as in G. laurifolia. This is net mentioned in the Flora Australiensis.

Spikes: Not truly secund at first, when young there are bracts all up the
rhachis (or all round). That is on the back, or the inner side not exposed and
facing the stem. As the flowers expand, they reach out towards the light; but
even then there are bracts on the back.

1This is what A. Cunningham means when he says of this species that its habitat is
“wet, peaty bogs on the Blue Mountains.”

BY C. T. MUSSON AND J. J. FLETCHER. 129

Flowers: In very young spikes, before the styles protrude, the flowers, like
the exposed surface of the bracts and the rhachis, are silky white, due to the
hairs, and show no colour, or very little. The bracts are greenish or yellowish
green on their inner surfaces. The bowed styles are rich pink; before they appear
a little colour shows in the perianth, but it is masked by the silky hairs. In older
spikes, it is to the massed styles the colour of the spikes is chiefly due, as seen
from a distance, a fact not usually recognized. As the flowers mature, the colour
of the styles fades to a lighter pink. At any time, viewed from outside, the
perianths show but little colour. But when the styles are nearly ready to
straighten, and afterwards, the inside of the perianth, viewed from front or above,
is a rich purple (sometimes looking almost black), fading to »yurplish red in
dry (herbarium) specimens. It fades considerably and what is left appears as
longitudinal streaks (about ten). In herbarium specimens the colour of the inside
of the perianth is perhaps more noticeable than in the fresh state; possibly due
to disarrangement and flattening of the silky hairs in the process of drying. The
perianth limb looks greenish outside, notwithstanding the coating of white silky
hairs, this silky indumentum being most conspicuous. The hairs are appressed,
projecting and rather tufty on the revolute limb; after the stigmatic disc has
been released they appear as a very noticeable tuft on each side of the expanded
revolute limb. The tube of the perianth of young flowers in bud, bracts and
rhachis all appear of the same colour when fresh. Flower pedicels are greenish
(like rhachis and bracts) and coated with white hairs, when fresh. The pollen
masses when freshly exposed on the stigmatic surface are pink, those of G. lauri-
folia and the hybrid (G. Gaudichaudii) are bright yellow. The stigmatic discs
are light green, fading to yellowish green. ;

ni
The Hybrids (Plate vii, figs. 2-5). Fresh flowers November, 1913.

With the exception of one plant, I have not seen spikes and flowers of any
of the Gaudichaudii series, that were not readily distinguishable from those of
acanthifolia. But I have had the great advantage of examining fresh as well as
dried flowers. Plants of Section i have flowers and spikes like G. laurifolia, a little
lighter in colour in some, but still some shade of crimson, with the same notice-
able contrast between the tube and the globular limbs of the perianth (crimson as
compared with ferruginous), but with a noticeable increase in the white silky
hairs on the tube giving this a sericeous appearance. The inner surface when
exposed purple. Plants of Section ii show a good deal of difference in the amount
of colour in the perianth, some having very little (not more than in acanthifolia)
when growing covered up in the shade, except as regards the purple inner surface,
when the styles are hardly darker than those of acanthifolia. In others there is
more diffused colour, lighter than in laurifolia, the exposed inner surface purple,
when the limb and tube split. The pedicels are green as in acanthifolia, but
what readily distinguishes them from that species is the admixture of white and
crimson hairs on the limb, and white and ferruginous hairs on the globular end.
The tomentum as a whole is also more copious and longer on the globular end.
Even in dried specimens the contrast is strikingly obvious. The rhachis and
bracts are also distinguishable from their rusty appearance due to the admixture
of white and rusty or ferruginous hairs. This admixture of the different kinds
of hairs on the flowers, rhachides and bracts is perhaps one of the best examples
of mixed characters in these hybrids; apparently analogous to the case of hybrid
roses between parents having either glandular or non-glandular hairs, mentioned

Q

130 NATURAL HYBRIDISM IN GENUS GREVILLEFA.

by Kerner.1 The tomentum, as a whole, is more copious than in laurifolia, but ~
less so than in acanthifolia; the hairs are somewhat longer than in laurifolia,
not quite so long as in acanthifolia. The hairs on the ovary are more numerous,
and the tuft is denser than in lauwrifolia, being very -like acanthifolia, but in
many cases there are a few crimson hairs mixed with the white.

Classification of Hybrids (G. Gaudichaudii).

The series of hybrids met with, comprises recognizably different types of
individual plants. The series, as known to me (J.J.F.) is divisible into two
sections according as the plants have entire mixed with pinnatifid leaves, or all
leaves are pinnatifid, as in G. acanthifolia the number of leaf lobes is not constant.
With this difference in leaf character there are correlated certain differences in
flower character.

Fresh flowers are only available during the summer months, whereas plants
may be seen in leaf through the year, therefore the leaf characters most readily
catch the eye. I have had specimens, illustrating a series of stages commencing
with laurifolia and ending near acanthifolia, whilst the reverse could also, neces-
sarily, be illustrated. A short account of various types will show how the
characters of the parent species are blended, also indicating something of the
variations to be seen, pointing conclusively to their hybrid nature.

Section i: Prostrate plants with procumbent stems. Entire leaves altogether
of the laurifolia type, mixed with pinnatifid leaves with lobes numbering up to
9, all the lobes entire. Flowers indistinguishable from laurifolia, or with more
white appressed hairs on the tube of the perianth. Habit like that of laurifolia.

A. Entire leaves most numerous with pinnatifid leaves usually having not
more than three-lobes (very rarely four). Flowers and inflorescence
indistinguishable from those of laurifolia.

B. Entire leaves in a minority, pinnatifid leaves, up to five lobes; venation
and indumentum of laurifolia; tube of perianth with more white hairs.

C. Like B, but indumentum reduced to'a remnant of scattered appressed
hairs occurring singly, so that underside of leaves is almost glabrous,
much as in some plants of A.

D. Entire leaves more reduced in number mixed with pinnatifid leaves,
having 2-7 or any intermediate even number of lobes, indumentum well
developed in some specimens.

Section ii: Prostrate plants with procumbent stems, stiffer than in laurifolia,
and not forming such leafy carpets. All leaves pinnatifid, with from 3-15 lobes
(maximum and minimum not found on same plant). Some leaves with entire lobes,
but in every plant seen, some bilobed. Three and four lobed lobes may occur,
occasionally with bilobed on the same leaf. Indumentum very variable, well

1 The cellular structures produced from the epidermis of the stem and leaves which
are differentiated as hairs, bristles, scales, glands et cetera, classed together as under
the name of indumentum, are very constant characters in most species of plants.
Hybrids exhibit the most varied combinations of the indumenta of their parents. In the
majority of cases the characteristics of the two stocks in this respect are mixed, but less
frequently they are united, and in the latter case the shape, size and number of hairs
tra are intermediate between those of the appendages in the two parent species.’’—
Kerner and Oliver, Nat. Hist. of Plants, p. 564.

“Where one parent rose bears only non-glandular and the other only glandular hairs,
the hybrid is sure to be clothed with a mixture of the two kinds of hairs.’’—Ibid., p. 564.

“The colour of the flowers in hybrids is usually the result of a fusion of the colours
in the parent species; less frequently it is a mixture of the original colours.’—Jbid., p. 567.

BY C. T. MUSSON AND J. J. FLETCHER. ~ Bil

developed and much reduced on the same plant. Intermediate in extent on
others, or reduced generally, on the leaves to a functionless remnant of appressed
hairs, singly scattered; recognizable but vestigial. No plant seen with under
surface of leaves entirely glabrous.

Fresh flowers with more white silky hairs on tube of perianth, hiding the
colour and giving dried flowers a greyish appearance. On the revolute limb of
the perianth the hairs are tinged with ferruginous or reddish colour. Capsules
hardly distinguishable from those of acanthifolia.

E. Leaves with from 3-11 lobes, the maximum of indumentum for this
section, but it varies in amount on leaves of the same plant.

F. (Here is G. Gaudichaudii R.Br.). Leaves with from 3-11 or 5-15 lobes,
with a maximum of bilobing; indumentum recognizable but vestigial,
reduced to a few scattered hairs. Perianth with the tube sericeous, the
hairs on the revolute limb tinged with ferruginous or reddish. This is
the type of hybrid most frequently met with.

G. Leaves with 7-13 lobes (or any intermediate even number), with most
(but not all) leaves having a maximum of secondary lobing; some tri-
lobed, rarely four lobed. No five lobed lobes seen.

Remarks on the Various Types of the Series.

A. If the pinnatifid leaves are removed, it will pass as a specimen of
G. laurifolia. If it were the only member of the series Known, it might be
regarded as a sport, perhaps due to bud variation. Fine colonies of G. laurifolia
carpeting considerable areas of ground may be found too remote from plants of
G. acanthifolia for birds to pass directly from one to the other, but in such cases
the plants show no tendency whatever to produce lobed leaves. Fourteen plants .
of this type seen; from the conditions under which they were growing, I believe
them to be seedlings from ovules of G. laurifolia fertilized by pollen of
G. acanthifolia.

One fine plant was growing in the midst of a carpet of G. laurifolia, and, except
in one instance, the others were growing quite close to, side by side with and the
branches overlapping, or a little lower down a slope than, one or more plants of
G. laurifolia. In some cases they were the only hybrids to be found in the
locality. In one case only was a plant solitary, and this must have been a case
either of a seed having been removed further than usual from the parent form, or
more probably was a plant whose former associates had been removed by fire or
accident.

B. Six plants have been examined. The finest example was growing on a
grassy slope, well exposed to the sun, just below, several plants of G. laurifolia
with plants of G. acanthifolia a few yards away at the bottom of the slope. Another
was one of a row of four contiguous plants; the two middle ones were examples
of type C. The remaining one of the four a fine example of type F. Two others
were growing close to plants of G. acanthifolia and the sixth quite close to a group
of plants of that species.

C. Two plants only seen, the two referred to above. Both old, very dry and
quite alike. Dried leaves appear almost glabrous, but the scattered hairs readily
seen on fresh leaves. They had a very distinctive appearance. The leaves, even
when fresh, were incurved at the edges in an unusual manner. The facies of
these two plants suggested the idea that they were suffering from excessive trans-
piration due to the loss of the indumentum.

132 NATURAL HYBRIDISM IN GENUS GREVILLEA.

D. Three plants seen, all, I believe, seedlings from G. laurifolia. One was
growing in the midst of a carpet of G. laurifolia, another was beside a plant of
that species, the branches overlapping. The third was growing in a grassy glade,
near some other hybrids, but with plants of both G. laurifolia and G. acanthifolia
close at hand.

BE. Two plants seen, both remarkable for the well developed indumentum
and for its variable amount. One plant showed great variety of lobing in the
leaves. The other was overshadowed by a plant of G. acanthifolia, and hampered
in symmetrical growth by other plants. A well marked intra-marginal vein
present. But for the indumentum, this plant was quite of the F type.

F (= G. Gaudichaudii, R.Br.). The commonest type met with, about fifty
plants seen. Not only is this the type most frequently produced, but in my opinion it
is the reciprocal hybrid. Solitary plants were seen in the midst of carpets of
G. laurifolia; others were seen growing close to, even overshadowed by, a plant
of G. acanthifolia. Others were growing on slopes between G. laurifolia above and
G. acanthifolia below. On the whole they are most frequently situated near plants
of G. laurifolia. Mr. Forsyth noted this close association with G. laurifolia, but
without realizing its significance. The reason, as one can see, is that when the
birds: travel up the valleys visiting G. acanthifolia in the swampy lower parts first,
and come to the last of them, they very naturally visit G. laurifolia on the slopes
above, if there is a display within sight. On the other hand, after visiting the
Banksias, Lambertias, and other plants on the ridges and upper slopes they work
down over the laurifolia areas until they reach the wetter lower slopes where
acanthifolia flourishes; they may then pass direct from the former species to the
latter.

G. Five plants seen, two of them fine plants, one with stems seven feet long,
growing in the midst of a carpet of G. laurifolia; another near a plant of type B,
close to plants of both parent species. A fourth was growing between plants of
the two parents. The fifth, a solitary plant, growing further from the parent
forms than usual; but there was evidence of interference with the surface.

Summary.

Grevillea Gaudichaudii, R. Br. is a hybrid. The parents are G. laurifolia Sieb.
and G. acanthifolia, A.C.

The forms described by R. Brown, Gaudichaud and Bentham are representatives
of naturally related forms, the result of hybridization; and fill places in a graded
series between the parents.

These plants are only found near one or both parents, along the Blue
Mountains from Wentworth Falls to Blackheath. (The area west of this has not
been searched.)

The characters of G. Gaudichaudii, R. Br. are blends or mixtures of those of
the parent forms.—C.T.M. 2

EXPLANATION OF PLATE VIL.
1. Grevillea laurifolia, Sieb.
2-5. Various forms of Grevillea Gaudichaudii.
6. Grevillea acanthifolia, A.C.

1The Spinebill chiefly.

THE PHYSIOGRAPHIC AND CLIMATIC FACTORS CONTROLLING THE
FLOODING OF THE HAWKESBURY RIVER AT WINDSOR.

By Lestry D. Hatt, B.Se., Science Research Scholar in Geography,
the University of Sydney.
(From the Department of Geography.)

(Plate viii, and nine Text-figures. )
[Read 27th April, 1927.]

Introduction.

Floods of the Hawkesbury River at Windsor have been part cf man’s experi-
ence ever since the first farmers came and settled the rich alluvial flats along its
banks in 1794. Records show that in recent years these floods have decreased
considerably both in height and frequency, but the cause of the decrease is not
definitely known. It may be due either to dam building on the headwaters of
the river, or to an actual diminution in precipitation. In the following paper the
effect of the Nepean dams on the floods of May and June, 1925, has been investi-
gated in an effort to find a definite relationship between flood waters, water
conservation and climatic conditions over the catchment area of the river. The
term “catchment area” is used throughout to denote the entire area drained ‘by
the stream irrespective of special regions reserved for water supply.

So far as I am aware no work of this nature has been done for the Windsor
district. Josephson, 1885, discusses the history of the floods of the Hawkesbury
River, but makes no attempt to give their causes or to enter into the details of
rainfall. David and Guthrie (1904), have investigated the nature of the Hawkes-
bury River silts and their results are mentioned in this work. Information as to
the position and capacity of the dams and the amount of water stored during
the flood months, were kindly supplied by the Metropolitan Board of Water Supply
and Sewerage; while other details concerning rainfall and run-off for the whole
Windsor catchment area were contributed by the Water Conservation and Irriga-
tion Commission. Thanks are especially due to Mr. French of this Commission
for making these facts available and for his helpful suggestions and kindly
assistance. Rainfall figures were given by the Weather Bureau and flood details
were obtained from copies of the Windsor and Richmond Gazette of various dates,
for which my thanks are due to the Editor who gave me access to his files.

GENERAL DESCRIPTION OF THE HAWKESBURY RIVER.

Three hundred and thirty-five miles in length, the river with its several large
tributaries, drains all the immediate country to the north, west and south of
Sydney, an area of approximately 8,000 square miles. It rises 20 miles from
Goulburn. where it is known as the Wollondilly, flows generally northwards, and
after receiving water from the Cookbundoon, Wingecarribee and Nattai streams,

is Joined by Cox River from the western mountain region. From here it is known
R

FLOODING OF HAWKESBURY RIVER AT WINDSOR.

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as the Warragamba and flows through a deep gorge till joined by the Nepean near
Mulgoa. The main river is now known as the Nepean and continuing northwards,
is joined by the Grose a little above Richmond and by South Creek at Windsor.
After the Grose junction it is called the Hawkesbury and is joined by two large
streams, the Colo and the Macdonald, before reaching Wiseman’s elbow, where it
turns eastward and finally enters the sea at the beautiful drowned estuary of
Broken Bay. A consideration of the map, Fig. 1, will show that the headwaters of
the river above Windsor may be divided into three systems, namely, the
Wollondilly-Cox, the Nepean and the Grose, each of which has its own definite
catchment area.

The Wollondilly-Cox has a catchment area of 3,383 square miles including all
the country drained by the Wollondilly, Cookbundoon, Wingecarribee, Nattai, Cox
and Warragamba rivers. Part of the western warp, it is all highland, but varies

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of the Hawkesbury River above Windsor.

somewhat geologically. The upper Wollondilly flows through a series of Palaeozoic
rocks, so that its gorges are less striking than the huge canyons which have been
cut in the Hawkesbury sandstone by the Cox. The lower Wollondilly, Nattai and
Warragamba have also cut deep gorges in the sandstone. Thus the northern part
of the catchment area is of a very rugged character making communication
difficult. Within the area there is a comparatively small population and only a few
large towns. Settlement has collected in three regions. On the Upper Wollondilly,
beyond the limits of the sandstone, there is good agricultural land which supports
a farming population. Goulburn, population 11,950, is an important inland town
on the Great Southern Railway. Taralga is another large town in this district
where grazing, dairy-farming and fruit growing are the chief industries. Good
workable soil and a less rugged topography, have led to settlement in this area.

136 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

In the Bundanoon-Mittagong district is a series of small towns along the
Southern Railway, well within the sandstone area and important only from the
tourist and residential aspect. Among these are Bowral (2,640), Moss Vale
(2,030), and Mittagong (1,440). Round Sutton Forest is some good grazing land
due to the presence of basalt and the intrusive rock of the Gib at Bowral is
important commercially. On the headwaters of the Cox are several towns the most
important Of which is Lithgow (12,940). Here river erosion has caused the sand-
stone to be worn away and the coal measure series to be exposed. Coal is mined
at Lithgow and is the cause of the industrial prosperity of that centre. Other
towns on the Cox catchment area are Mount Victoria, Katoomba and Wentworth
Falls, all of which are tourist resorts. Except for these regions, juvenile dis-
section and barren sandstone soils have prohibited settlement on the Warragamba
catchment area.

The Nepean.—On the headwaters of this stream 354 square miles of unin-
habited country have been reserved for Sydney water supply. The catchment
area is practically all Hawkesbury sandstone drained by the Nepean, Avon,
Cordeaux and Cataract rivers. The maximum elevation is only 2,000 feet, so the
river gorges are by no means as rugged as those of the western warp. Still the
district does not invite close settlement and the small towns are mostly confined
to the southern railway line. The presence of juvenile gorges at the headwaters
of the streams and the absence of any close settlement were two factors in
determining the choice of this site for the location of the water supply dams.

The Grose has a catchment area of 250 square miles; a sandstone region of
juvenile dissection and part of the western warp. Its deep canyons, wide gullies
and barren hill tops prohibit all settlement except for towns such as Springwood,
Lawson and Blackheath along the western railway, which are important tourist
resorts.

The waters from these three highland areas are discharged into a com-
paratively flat region of Wianamatta Shale and lake silts; undulating: grassy
country with hills not more than 200 feet above sea-level, which contrasts strongly
with the sandstone highlands surrounding it on three sides (Taylor, 1923a). The
Nepean leaves the southern warp at Menangle and meanders through the “fossil
lakes” of Camden and Wallacia before again entering the uplifted region where
it joins the Warragamba. The Camden district, therefore, is only liable to be
flooded by Nepean water, which comes from a smaller drainage area than that
which affects Windsor. The combined Warragamba and Nepean water is dis-
charged on to the flats again at Penrith, causing floods here before it is joined
by the waters of the Grose. The floods on the lowlands of Richmond and Windsor
are caused by waters draining a total catchment area of approximately 4,840
square miles. At Cattai the river enters the northern warp and the flood waters
are once more contained within the river valley.

History oF Froops at WINDSOR.

The Hawkesbury River is recorded to have flooded for the first time in 1799,
doing much damage and causing great distress among the people of the young
settlement. This flood ushered in a series from 1799 to 1801 and then the river
was normal till 1805 when another series of floods commenced. The records of
these early floods are not accurate and no attempt was made to ascertain to what
height the river rose above normal. In 1808 there was another flood and the
frequency and destructive power of these floods was the cause of much anxiety

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138 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

to the infant colony. Thus we find that the town of Windsor owes its origin
directly to the necessity of constructing large granaries at higher levels in order to
protect the produce from inundation. During 1811 a severe drought was experienced
and was followed directly by another flood. Situated under the shadow of the
great warp, the Windsor district does not receive a very high rainfall and
droughts are fairly frequent, while the rain falling on the mountains will often
cause floods of the river. These unsuitable conditions and the growth of the
settlement made it imperative that the mountains should be crossed and more
extensive pasture lands discovered. Between 1816 and 1819 three large floods
occurred, then we have record of a series of smaller floods occurring at intervals
until 1857 which marks the beginning of a wetter cycle. Three floods are recorded
for 1860 followed by a small one in 1863 and a very large one in 1864. Three years

SSE

= —S=

TES eg cena op 3 Sane a4 ,
ilbers } Ene eros gees © 1 See Se
BEES An on

6 a = =
ingula “-——~e=

Text-figure 4.—Windsor Peninsula surrounded by flood waters, June, 1925.
(Sketch from a Daily Guardian photograph.)

later came the largest flood known to white men when the water rose 63 feet
above summer level of the river, entered the town and left only two parts of it
exposed as islands.

The people had hardly recovered from this disaster when, in 1870 the river
rose 15 times and flooded the flats five times. This was a record wet year at
Windsor and the climax of the rainfall which had been increasing since 1857.
From 1870 to 1879 floods occurred of smaller magnitude than the preceding ones,
after which a drought period set in so that the river did not flood again till 1889. ,
For the next few years a good rainfall was recorded, but at the close of 1893 a
drought began which lasted about fourteen years, although there were floods at
Windsor almost every year between 1893 and 1900. Since that time the rainfall
has been steadily decreasing and though some floods have occurred their height
has been much less. In 1904 the river rose over 40 feet, a record which has not
been equalled since. Dry years followed in which the rainfall for the year was
below the average until 1910, when it started to increase and a small flood of
21 feet occurred. 1911, 1912 and 1913 had small floods without much rain in

BY LESLEY D. HALL. 139

the Windsor district, then in 1915 heavy rainfall throughout the Hawkesbury
watershed caused a rise in the river of 25 feet. October, 1916, witnessed a high
flood of 35 feet 6 inches, followed by five years of drought which were broken by
heavy rains in 1921, when the river rose 20 feet. From this time onwards the
rainfall has been better, the flood of 1922 being a fairly high one. There is no
doubt, however, that floods of the Hawkesbury have decreased considerably during
the last twenty-five years. .

In May, 1925, the water rose 25 feet, that is, about 2 feet over the Windsor
Bridge, and a great many of the low-lying places were covered. The water only
broke over the banks of the river in very low spots and, running up the gullies,
inundated a good deal of land. In the flood of June the water was 15 feet above
Windsor Bridge, or 37 feet 3 inches above summer level of the river, which is the
biggest flood experienced since 1904. Many people on the lowlands were driven
from their homes and it is estimated that damage amounting to between £40,000
and £50,000 was done in the Windsor district and approximately £30,000 round
Richmond. All the country round Cornwallis, Freeman’s Reach, Pitt Town
Bottoms, Wilberforce lowland, the Peninsula and the flats along South Creek
were one vast sheet of water dotted here and there with the tops of houses and
trees. Windsor was a complete island as the water backed up over the roads at
the southern end of the town. The following is a list of the floods which occurred
between 1795 and 1925 compiled from various sources:

Height Height | Height
above a above | above
Year. Month. summer Year. Month. summer Year. Month. summer
level, feet. | level, feet. level, feet.
1795 Jan. — 1857 Aug. 372 1889 May 383
1795 Aug. — 1860 Feb. 2632 1890 Mar. 38%
1799 Mar. 50 1860 Apr. 362 1890 Mar. 34k
1800 Mar. — 1860 July 342 1890 June 223
1801 Mar. — 1860 Nov. | 36 1891 June | 353
1805 Feb. — 1861 Apr. 274 nW23 a 123 ©) 6 263%
1805 Mar. — 1863 — — 1893 | Mar. 283
1806 Mar. 48 1864 June 48 1894 Mar. 318
1806 Cet. 3 1864 June 324% 1895 Jan. 302
1808 Nov. 20 1864 July | 36 1897 | Suly 22%
1809 May 48 1866 June 26 1898 Feb. 313
1809 Aug. 474 1866 | July 274 1899 Aug. 22
1810 July - 1867 Apr. 2 1899 Aug. 263
1811 Mar. — 1867 June 63 1900 July 46
1812 Mar. — 1869 May 362 1904 July 40
1816 June +52 1870 Apr. 45 1908 Aug. 15
1817 Feb. — 1870 May 358 1910 July. 21
1819 June 46 1871 May | 363 1911 Jan. 25%
1820 June — 1872 — = 1912 July 22
1821 Sept. — 1873 Feb. 413 1912 Aug. 24
1826 Jan. — 1875 June 38% 1913 May 264
1826 - Sept. — 1876 Sept. = 1913 May 20
1830 Apr. — 1877 May 30 1915 Jan. 25
1831 Apr. — 1877 May 20 1916 Oct. 354
1832 Mar. -= 1878 Feb. 262 TOI elle =
1833 | Apr. — 1879 | Sept. 433 1922 | July 31
1857 | July | 32h 1879 Sept. 342 1925 | May 25
1925 June 374

140 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

This list of floods records all those mentioned in the following references:

(1) 1799-1819: Articles in the Nepean Times for June, 1925, entitled ‘126
Years Ago.”

(2) Tebbutt: Catalogue of the floods and freshes of the Hawkesbury River
and South Creek at the Peninsula, Windsor, New South Wales. 1855-1903.

(3) “Results of Meteorological Observations at Mr. Tebbutt’s Observatory.
1898-1915.”

(4) Results of Rain and River Observations. 1903-1908.

(5) Windsor and Richmond Gazette. 1905-1925.

(6) Josephson, J. P., 1885: ‘‘History of Floods in the Hawkesbury River.”
Proc. Roy. Soc: N.S.W., page 97.

RELATION OF FLOOD HEIGHTS TO YEARLY RAINFALL.

Floods in the Windsor district come at all seasons and have been known to
occur in almost every month although the winter months, May, June and July,
are the most common (see graph, Text-fig. 5). No flood has been recorded for

7

a

Floocls.

S

u

Number of

O
Jan. Feb. Mar. Apr May June. July. Aug.

Month OF Year.

ept. Oct Nov. Dee. Jan.

Text-figure 5.—A graph showing the number of floods occurring in each
month of the year from 1795 to 1925.

December and very few in October or November. Floods are caused by heavy
Tain falling on the watershed of the river and are not affected by precipitation
at Windsor. A fall of rain on the southern drainage area sufficient to cause floods
is usually due to a cyclonic disturbance whose influence is felt all along the coast;
thus it often has rained or is raining at Windsor during the rise of the flood
waters. Cases have been known, however, when the river rose without any rain
having fallen in the immediate district. The graph, Text-figure 6, shows the floods
at Windsor in order of decreasing magnitude, with rainfall at Windsor for the
corresponding years superimposed. It is obvious that the two lines bear no
relation to each other. The third line shows the rainfall over the Nepean and
Warragamba catchment areas, calculated as an average from the four stations,
Goulburn, Taralga, Cordeaux and Picton. In this the relationship is more apparent
as the rainfall curve is composed of a series of maxima and minima, with a general
decrease in the height of the maxima, which corresponds to the decrease in height
of the floods. The minima mark those years when the heavy rainfall was more
coastal and all the flood waters came from the Nepean. The association between
the rainfall and the flood line is not very close, for a flood does not depend on

BY LESLEY D. HALL. 141

the amount of annual rainfall, but on a very heavy fall over a short period.
Constant precipitation may give a high annual total with not enough at any one
time to cause a flood, while flood rains followed by droughty conditions will
result in a low yearly average. If heavy rain falls at a time when evaporation
and percolation factors are high the run-off may not be sufficient to cause a flood.

WATER CONSERVATION ON THE NEPEAN.

The great flood of 1867 occurred during a time when floods were much more
frequent than they are at present and when the waters over the whole catchment
area were uncontrolled. Since that time schemes for the conservation of water
have been carried out on the Upper Nepean. The locality was chosen as being
the place nearest to Sydney which was high enough and comparatively unin-
habited; not from any direct rainfall considerations. The first project was the
building of a tunnel from Pheasant’s Nest to Broughton’s Pass, which diverted
water from the Nepean and Cordeaux rivers to the Cataract. The water of both
streams was carried along, partly by tunnel and partly in an open canal to

bo

50.

50

Floods at Wendsor, in order of greatest
magnitude. i

Rain at Windsor.
Rain on the Nepean and Cordeaux Catchment

20

Height of Floods in feet.

(average of % Stations, Goulburn, Taralga, Cordeaux, Reton
40 . 5
RIGO HWY KG eWY CSN HBS Qh HM so Ww: x = gi?
SONS SSR eee SONS Ses SSeS eee IRS
~m & ®& © % Gg & mn & BS OH © & & DH & & wm BH & OWweewo Qa
= oS SS Sp SNS SSS OO Sas Se SS Sse eS SS SS SS ST OS Seo ee

Text-figure 6.—A graph showing the relation of flood heights at Windsor to the
yearly rainfall. In this figure the continuous graph represents the floods at Windsor,
the dot and dash graph the rain at Windsor and the broken line the rain on the
Nepean and Cordeaux Catchments.

142 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

Prospect and so to Sydney. Then the Cataract dam was built and stored water
first in 1906. It was filled for the first time in 1911. In time of flood or fresh
only a fraction of the flow of the Nepean River can be diverted to Sydney and
the same might have been said of the Cataract River before the completion of
the dam. In dry weather, however, practically the whole flow of the Cordeaux

om

x Beton Bargo.
: a upton,

ams,

Pwversiggs

$5655.55 54 Ms a _
wee -"No. I
Se oan ane, a )
Cat ae
QW ne Dam
‘ @

CATCH MEN AREA. S

@ Letter ox
ae Sherbroa

EAE

Text-figure 7.—Sketch map of the catchment area of the Nepean River
showing the position of the water conservation dams.

above Pheasant’s Nest and the Cataract above Broughton’s Pass is diverted, to
the great disadvantage of settlers on the Nepean, who are thus deprived of water
which would have otherwise reached their land. To assist them compensa-
tion weirs have been built along the Nepean, and are replenished from time to
time by water released at Pheasant’s Nest and Broughton’s Pass.

The catchment area of the Cataract Dam is 50 square miles and, when full,
it backs up the water to a depth of 150 feet and stores 20,743 million gallons.
Construction of dams on the Cordeaux and Avon rivers is now almost complete
and during the flood rains of May and June, 1925, they stored a great deal of

BY LESLEY D. HALL. 143

water. As soon as these dams are finished one is to be constructed at Lupton
on the Nepean proper, and with the four dams in working order a continuous flow
of 108 to 110 million gallons of water per day will be able to be maintained to
Sydney as well as the coastal supply to Mt. Kembla and Wollongong. In 1871
there were only two rainfall observing stations in the catchment area but at
various times since then others have been established. The following is a list
of these recording stations:

|
Catchment Area. Rain Gauge Stations. No. of Years. Yearly Average.
Sherbrookciaeananeeeeeae 32 59-16
Madden’s' Plains) 2.5... | 17 58-47
Cataract. etter boxe. Secale oars tenes 16 49-43
Mount ebleasaniteeorycriarnK 174 57-13
Cataracty anime wrrieeeers 20 34-82
Broughtonispeassi esse 36 33-16
COndeauxe Weyer sic shessty cuss 53 61-01
Cordeaux Gauge Weir .. 15 42-04
Nepean, TROMSESIMENOD, | “Gaoococcsuo0eed 34 57-69
Mattar om Sgn ape tecsusroton! sieve aac 28 33-61
Bar eo Mrarsspses io ere Sere sors ols 22 30-22
WV BOTIE seer sewer cceteh cts ronsene shes 28 27:23

The average rainfall on the Cataract catchment area of 50 square miles
is 48-69 inches. On the Nepean catchment area, which includes 35 square miles
of Cordeaux drainage and-55 square miles of the Avon, the rainfall is less, the
average being only 41:98 inches. Therefore, though the Cataract Dam has a smaller
catchment area than the Avon it receives a greater amount of water. One inch
of rain falling over one square mile of country is equivalent to 14:52 million
gallons of water. A great deal of this water is lost by percolation and evaporation
so that only a small percentage runs over the surface and into the river. Measure-
ments of the amount of rain falling on the catchment area of a river will give
the actual number of gallons which fell, while from measurements of the velocity
of the stream the number of gallons discharged can be obtained. From these two
sets of figures the actual amount of water lost by percolation and evaporation can
be found and the run-off expressed as a percentage of the rainfall. Between 1887
and 1906 on the Warragamba catchment area the average rainfall was 32:45
inches and the mean estimated run-off was 16%. The run-off is found, to be a
variable factor depending on climatic conditions and whether the rain falls after
a wet or a dry season. On the Cataract catchment area for the 25 years since
1900, the average rainfall per annum was 50-56 inches and the average discharge
was 13,358 million gallons or 33-64%. An inch of'rain over the 50 square miles
of the Cataract catchment area is equivalent to 50 x 14:52 x 10° or 726 million
gallons of water. Of this about one-third or 242 million gallons is available to be
stored in the dam. The average fall of 50 inches of rain over the whole area will
only produce 12,100 million gallons of water to be stored and, as this is only
about half the capacity of the dam it seems evident that its catchment area is not

144 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

large enough. Yet the Avon Dam with about equal available water is twice as
big as Cataract with a total capacity of 47,159 million gallons. The heavy rains
of May and June, 1925, were not sufficient to fill the Avon and Cordeaux dams
which would seem to be too large for their catchment areas. In this connection
it is necessary to remember the purity of the water which flows into the Avon Dam,
an important factor in a country where settlements are to be found on most of
the rivers.

Tue ACTION OF THE DAMS DURING THE 1925 Froops.

Heavy rains were experienced on the Nepean catchment area during May and
June, 1925, the total precipitation over the whole area for the two months, as
ealeulated from four rainfall stations on the Cataract catchment and three on
the Avon and Cordeaux, being 37:08 inches. Of this 22-84 inches fell during May
as compared with the 14:24 inches which fell in June. At the time of the May
fall, however, the ground was dry and a great deal of the water was lost by
percolation and evaporation. For this month the average run-off over the whole
area was 63:°6% of the rainfall, while during June when the ground was still moist
the average run-off was 76:9%. On the Cataract catchment area the mean precipi-
tation for the month of May was 25:24 inches and the run-off was 66% of the rain-
fall. There were 5,470 million gallons of water stored at the beginning of the
rain and the increase was 6,263 million gallons. The dam did not overfiow during
the May flood and the water continued to rise during the rest of the month owing
to run-off from the hills which continues for a good while after rain has fallen.
At the beginning of the heavy June rains there were 17,954 million gallons in
storage and the dam was filled within three days. The overflow at the time of
the June flood is estimated at 4,421,324,500 gallons. The rainfall for that month
was 13:95 inches and the run-off was 74:7%. The total amount of water stored
in the Cataract dam in May and June was 15,273,000,000 gallons.

On the Cordeaux catchment area records from three stations only are known
and their readings give a mean rainfall of 23:19 inches for May. The increase in
storage of the dam was 6,848,000,000 gallons and the run-off was 58-4% of the
rainfall. The June rains averaged.15:29 inches but the run-off was higher, 70:3%,
and increased the storage by 5,439,000,000 gallons, making a total increase of
12,287,000,000 gallons in the Cordeaux storage for May and June. The dam did
not reach its full capacity. Records from the Avon catchment area are also very
approximate, being based on observations from three stations only. For May a
mean precipitation of 20-1 inches is recorded, 25% less than that of Cataract,
while the run-off was 66% and gave an increase in the storage of 10,625 million
gallons. In June the rainfall of 13:47 inches with a high run-off of 85:8% gave
an increase in storage of 9,193,000,000 gallons. The total increase in the Avon
storage was 19,816,000,000 gallons, but as the dam was almost empty at the begin-
ning of the rains it did not then reach its full capacity. Storage was commenced
on the Avon River at the end of May, 1924, and by the end of June, 1925, 22,308
million gallons had been impounded; practically the whole yield at the catchment,
as only a trifling amount was sent to Sydney. Another 71 inches were then
required to fill the storage. It is calculated that the mean precipitation over the
whole catchment area of 354 square miles for May and June was 18-54 inches,
giving a run-off of 105,372 million gallons, of which 48,233,000,000 gallons, or
nearly 50% was caught and held in the Cataract, Avon and Cordeaux reservoirs.
Some of the excess water not held by the dams was probably discharged down

BY LESLEY D. HALL. 145

the canal to Prospect, but in quantity so small as not to affect the calculations,
since at the height of the flood the gates are closed to prevent the blocking of the
canal by debris.

Details as to the rainfall and discharge from the Warragamba and Grose
catchment areas are supplied by the following letter from the Water Conservation
and Irrigation Commission: “The most notable falls on the Warragamba River
catchment area during the months of May and June were:

Robertson LAT Mit Mndties, pe 44°20) inches
Goulburn SP ARERR A. te celOs38 Py
Katoomba Se See ae ye aad MAAS e
Wiha Wiiewore, oo co oc ao IbupmHy) h
Bowral te Sse ee i sscaes eeepc £9 ki
Taralga mabe. Tey a Ys ote Ss LOOP 6

The rainfall stations are not evenly distributed over the catchment area; on
the sparsely settled districts the records are scanty, but by giving each station its
proper ‘zone of influence’ the mean depth of precipitation over the catchment
area is estimated at 17 inches. The amount of water discharged by the Warra-
gamba at a point a litttle up stream from its junction with the Nepean, during May
and June is estimated at 297,715,000,000 gallons or enough to supply Sydney for 14
years at the present annual consumption rate. About 35:5% of the rainfall was
discharged by the Warragamba, the rest being lost by percolation and evaporation.
There are no reliable measurements of the amount of water contributed by the
Grose River to the floods of May and June. The catchment area of the stream at
its affluence with the Hawkesbury above Richmond is 250 square miles. A con-
servative estimate of the discharge, basing it on the relation of its area to that
of the Warragamba, would be 22,000 million gallons of water, which is little more
than the capacity of Cataract Dam. Rainfall stations on the Grose catchment
area are Springwood, Lawson and Blackheath.”’

From a combination of the figures for the three catchment areas the following
table is derived:

Area in Run-off, Storage,
Catchment. square million million Storage %.
miles. gallons. gallons.
Wiarracambayiesy wee fhe hee os 3,383 297,715 — “=
i
Nepean Seot eich eT Pete OR, ao Ont 354 105,372 48,233 45-7
Grose aigiyett ato o. Mesoraes Ghee Ie RRSnOaeteT 250 22,000 — —
PIN OVATE ee RP a ai | oie iat otal Bic 3,987 425,087 | 48,233 11:3

This gives approximately the total run-off from a catchment area of 3,987 square
miles, figures for the other 853 square miles of the Windsor catchment area not

being recorded. Yet considering that the dams caught and held almost half the
7

146 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

water discharged from the Nepean catchment area and practically 10% of the total
water discharged at Windsor, it is obvious that if these dams had not been in
existence the magnitude of the floods at Penrith and Windsor would have been
greatly increased. The dams on the Nepean cannot prevent floods at Windsor,
nor decrease the number experienced there, but their existence is felt in a tendency
towards modification during a flood period, although their effect is a variable factor
depending on a number of conditions. The Nepean drainage is only about 10%
of the Warragamba, but its area is nearer the coast and consequently receives
more rainfall, so that in the June, 1925 flood its run-off was equal to 35% of the
Warragamba run-off. It is possible for coastal rains to cause floods of the Nepean
without additional water from the Warragamba and in such cases there is no
doubt that the effect of the dams is very great. The actual amount of water which
is held back by the dams depends on the number of gallons already stored at the
beginning of the rains. The heavy rains of May were preceded by a dry period
and the water in Cataract Dam was low. Also dams at Cordeaux and Avon were
only on the verge of completion and almost empty, so that a considerable quantity
of water was capable of being stored.

A flood of any magnitude at Windsor is due to a cyclonic disturbance, which
causes sudden heavy rain of only short duration over the whole catchment area.
As a result a great deal of excess water is discharged in a few days, causing an
abrupt rise and fall in the river level. The dams, therefore, by holding back
water at the-.time of general discharge decrease the height to which the flood
would have risen and the consequent damage which would have been done. HEven
if the dams are full and cannot hold back the water from the flood they decrease
its peak discharge, because the whole surface of still water must be raised above
the level of the dam before the overflow takes place and the drainage from such
a surface is necessarily slower. It is not the actual amount of water but the rate
of discharge which causes the river to rise and flood the lowlands and an arrange-
ment whereby the water is delayed and caused to pass more slowly down the river
does much to lessen its evil effects.

DECREASE IN RAINFALL.

The decrease in the number of floods experienced at Windsor of late years
seems rather to be a factor of the decreasing rainfall than to be due to dam
building on the Nepean. At Grove Farm the river rose to 67 feet 10 inches on
the present gauge in 1867 and only 35 feet 4 inches in 1925. This difference could
not be accounted for by the building of the dams. It must be associated with a
diminution in rainfall, a factor not due to deforestation as this area, being rugged
inhospitable country, is still uncleared to a large extent. The difficulty in this
connection is the absence of long time rainfall records by which a decline in
precipitation might be verified. Goulburn is the oldest established rainfall station
on the Warragamba catchment area. The records commenced in 1858 and the
average for 67 years was 25-41 inches, while for the last 30 years the average has
been 22-83 inches, a falling off of about 10%. Rainfall records at Cordeaux
gauging weir were started in 1872, the average rainfall for 53 years being 53:58
inches, while the average since 1900 shows a decrease of about 20% and is only
44-45 inches. The following table shows how the ten yearly averages compare
with the total averages since the formation of the station at various places on the
catchment area (figures obtained from Hunt, 1916, and from the Weather Bureau).

ILALL.

BY LESLEY D.

(hOO0

'3oc

shva2ny

SS eee

Lithgow-
Me Viectorva
Actual Raunfall.
Average Ratnrall.

Nn —_
ba [?
/;
uv
>
fe)
Ho)
<

La
Kapeoey

300
200
100

3 8 8

‘qqeduivy do sayouy

1200

1100

1000
Joo
300
700
6

4ebI
6b!
trbl
Itbh
orb
bibt
8161
Ltbl
VItbt
Gibt
W161
1b
Tibet
11bt
orbl
bobl
Bobr
Lobi
1061
Sobt
4obl
G06!

Sa See ae a
SS MoT iobt

9994

Pees oS

SS? Gusany se? u
oe eA

figure 8.—Summation hydrographs showing the variation in the rainfall above

and below the average for a number of years, of six stations on the Windsor catchment

Text-
area.

The summation hydrograph is made by adding together the averages of the rainfall
at a certain place over a number of years and then plotting the result as a rising graph.
The final point on the graph is joined to the first point by a straight line which is the

general average rainfall over the total number of years.

When the graph line is parallel

Where it inclines away from

to the average line the years were ones of normal rainfall.

the average the rainfall during those years was above normal and where it inclines in to

the average the rainfall was less than normal.

148 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

Average Rainfall.
Total
IRIS: 1871— | gsteen) tsg1— || loots | tones ||) onsale eas
1880. | 1890. 1900. 1910. | 1920. 1924.
Cordeaux seer 60-0 | 57-42 69-66 46-34 47-91 39-58 53-48
|
ipictonuasee eee = 27-45 35-26 24-74 34-94 28-38 30-15
Maralleala eee = 30-87 30-35 24-49 28-69 27-05 28-28
Goulburn ...... 28-51 22-53 27-21 20-39 23-58 26-038 27-06
|

1871-1880 are years of good rainfall, 1881-1890 are not so good. From 1891-1900
the stations all record rainfall above the average, then from 1901-1910 the rainfall
is uniformly low. 1911-1920 shows an increase with two stations above and two
below, but 1921 to 1924 are years of low rainfall again. The rainfall at these
stations is also depicted on the graph (Text-figure 8), where summation hydro-
graphs show the variation above and below the average from year to year. These
graphs show that since 1900 the yearly rainfall has mostly been below the average
with a marked decrease in the number of inches precipitated. For six years, 1904-
1910, there was a low water condition in the upper Sydney water supply canal
from the Nepean, unparalleled in 80 years, and though the average rainfall
discharge has increased since then, it is still lower than it was before the dry
period.

The graph (Text-figure 9) shows the 11 year cycle of maximum and minimum
sun spots with a decrease in the maximum number from 1870 onwards. 1870 was
a year of good rainfall everywhere and a time when the greatest number of sun

Text-figure 9—Major and minor sun spot cycles (Huntington, 1915).

spots have been recorded. From 1881 to 1890 the rainfall was not so good, but at
that time the sun spot maximum was not nearly so large as the one preceding.
Between 1891 and 1900 the maximum was greater and the rainfall recorded is
better, but from that time onwards there has been a steady deciine both in sun
spots and precipitation. Thus rainfall does seem to vary, not only with the eleven
years’ cycle but possibly also in accordance with a superimposed major cycle as
shown by Huntington, 1915. If this is the case the recent floods of 1925 might be
taken to indicate the beginning of a new cycle in which the seasons would
gradually improve. Such conclusions are necessarily very tentative as it is
difficult to arrive at any definite principle.

BY LESLEY D. HALL. 149

FrLoop SILTS OF THE HAWKESBURY RIVER.

The silts brought down by the Hawkesbury when in flood are of two varieties,
the rich black mud and the useless sand. The Warragamba, Grose and Nepean
flow through sandstone country which is easily weathered, supplying these rivers
with large quantities of sandy sediment of no fertilizing value. The mud which is
so important to the farmer is derived by the Cox and Wollondilly from the older
palaeozoic rocks through which they flow before entering the sandstone region. By
the time the waters reach Windsor the sand and the muds have been mixed together
giving a light sandy loam which is very fertile. Pure mud is also deposited,
enriching the land, but sometimes the flood currents leave behind them patches of
sand which ruin the area on which they are placed. The Grose River carries only
sandy sediments yet after the flood of June, 1925, black mud inches deep was
deposited along the banks of this stream for seven miles above its junction with
the Nepean, thus showing the distance to which the flood water must have backed
up the Grose against the normal flow of the river. The depth of silt left behind by
the flood is a variable factor depending largely on the topography of the district,
but also on the height of the waters and their velocity. As the river spreads over
the flats it meets resistance and the velocity is checked causing a great deal of the
silt load to be immediately dropped. In this way levees or mounds of silt are
formed along the banks of the river which prevent the overflow water from drain-
ing away from the flats again. In the Windsor district, therefore, large areas of
still water result, the silts of which are deposited in a fine layer over the entire
surface. Farther down the river where the flats are smaller and more isolated
the silt deposit is greater and was from 4 to 6 inches in the June flood. As seen
in the river bank at Windsor the flood silts form a series of fine laminations of
varying colour according to the proportion of mud and sand (Plate viii, fig. 1). As
the surface layer dries it hardens and contracts, breaking into small pieces by a
number of small and irregular cracks (Plate viii, fig. 2). When dry it is very
soft and friable and soon becomes worn down into the soil. David and Guthrie
(1904) give an analysis of flood silt from the Hawkesbury water during the flood
of 1904 which shows that the lime, potash, phosphoric acid and nitrogen content
of the soil is renewed by flood waters. This is undoubtedly very good, especially
in a place such as the Hawkesbury flats where constant tilling is wearing out the
good of the soil and artificial manures must be used.

Provided the flood comes at the right season it is more beneficial than other-
wise in the more northerly reaches of the river where it is only the orchard flats
which are covered by the waters. Here the houses are built on the higher sand-
stone terraces and so escape damage. The fruit trees are grown in the long flat
gullies at the base of the sandstone hills where the flood forms quiet backwaters
in which silt is deposited inches deep over the farms. However, floods are very
uncertain in their coming and on the low flats between Wilberforce and Windsor
often large areas of growing crops are destroyed in addition to other extensive
damage which tends to outweigh any good effect. After the flood of June, 1925,
the water remained on the surface for days which, although impeding the work
of reconstruction, enabled the subsoil to become thoroughly wet. As a result it was
possible for the farmer to grow valuable areas of lucerne, a crop which cannot be
grown there under ordinary rainfall conditions as rain does not penetrate deeply
enough into the soil. Moreover, throughout the dry summer which followed the
crops flourished on water stored beneath the surface of the ground. Therefore,
although the general opinion is one of censure against the floods, there is always

150 FLOODING OF HAWKESBURY RIVER AT WINDSOR.

some benefit to be derived from them. In the Windsor district another difficulty

at flood times is the shifting of the land as the river increases the scope of its
meanders. The velocity of the flood waters gives them great erosive power and
as they swing round a meander the river bank is washed away on one side
while sand and debris are deposited on the other. During the June flood a land
slide occurred in which 200 yards of the road along the bank was completely
washed away. With every flood the channel of the river is altered, the meanders
becoming more and more curved and unless they are prevented by artificial
means the meanders will become cut off with resulting ox-bow lake formation.
This action of the river is a source of great difficulty in a region where the land is
divided into private farms whose owners lose large blocks of land with every flood.

FLoop PREDICTION AND CONTROL.

As floods in the Windsor district are due to irregular cyclonic disturbances on
the coast, to forecast their occurrence would be a matter of some difficulty. At
one time the people were caught by the water when quite unprepared, but they are
now warned by telephone communication which gives them about 24 hours in
which to evacuate. The warnings, however, are never very accurate and the
people do not know how high the water may be expected to come, nor will they
leave their homes till they are quite sure that it will be necessary. The best way
of controlling flood waters is to build levees along the river banks high enough
to prevent the water from coming over. in the Windsor district this would be an
enormous task owing to the great extent of flat land through which the river flows
in that locality. Moreover the expense of such an undertaking would not be
justified by the value of the land. It is doubtful whether any farmer round
Windsor would be willing to go to the expense of erecting a bank to protect his
property against the ravages of the river even if it did not do good as well as harm.
The risk of life does not even cause them to take precautions, for many still live
in houses well within the danger zone and do not even keep a boat when their
only means of escape at flood times is by water. The rescue work is carried on
by special boats which have to be sent to the afflicted areas from Sydney.

To control floods in the Windsor district it has been suggested to build a
dam in the Warragamba gorge to collect all the run-off from the Wollondilly water-
shed and permit of it being released in more or less regulated volumes. A wall 300
feet high in this gorge would impound the water for 25 miles up the Wollondilly
and 20 miles up the Cox. Such a dam might not actually prevent floods for the
Burrenjuck Dam, which captures nearly all the headwaters of the Murrumbidgee,
was unable to prevent flooding of that river in 1925. However, besides lessening
the discharge by an amount proportional to the number of gallons already in the
dam at the beginning of the rains, it would offer a very large surface of still
water over which the excess would have to pass and would thus make the floods
less intense and therefore lower in height. If the overflow water were to be
discharged down the river more slowly it might possibly be contained within the
river banks. The dam would give a continual flow of 200 million gallons per day
to Sydney. It is calculated by water supply experts that up to 1967 Sydney, with
the 80 million gallons from the Nepean, will only require 160 million gallons from
the Warragamba, and if the surplus is not used it will bank up and overflow.
This will decrease its usefulness in the prevention of floods and in order to use
the excess water schemes for the generation of electric power and for the irriga-
tion of the lowlands have been suggested. The water from the dam on the

BY LESLEY D. HALL. 151

Murrumbidgee is used to irrigate land which is naturally fertile and only requires
water to make it good for agriculture. It is well drained with uniform surface
levels which make redistribution easy, while the low rainfall insures a constant
demand for water. Conditions are different in the county of Cumberland. Here
there are 26,390 acres of good river land, 14,300 acres of second class river land
and the rest is Wianamatta shale, poor soil almost worthless for cultivation with
unequal levels which would make the application of water a costly process. 'The
area supports 778 persons per square mile, including the city of Sydney and only
98 per square mile if that town is excluded. Also the rainfall is a very variable
factor and the demand for irrigation water would fluctuate. Yet on the river
flats the dam would greatly decrease the possibility of floods and irrigation, though
costly, would increase the productivity of the land which is the closest agricultural
region to the city of Sydney. Nevertheless the proposal to dam the Warragamba is
not much favoured by people of the Windsor district, who depend on the river
for their water supply. The towns of Richmond and Windsor both obtain water
from the river by electric pumps and anything which would check or interfere
with the flow of the river or any of its tributaries upstream must seriously
jeopardize the supply from the Hawkesbury, especially in dry seasons when the
salt water works its way towards Windsor rendering the river water unfit for use.

SUMMARY AND CONCLUSIONS.

Floods in the Windsor district are due to the presence of a river with wide-
spread upland drainage, discharging its waters from the mountains on to the flat
lands of this locality. They are caused mainly by heavy coastal storms of short
duration and result from the sudden and simultaneous discharge of excess water
from the large number of tributary streams. This gives a typical high peaked
flood wave which passes very quickly so that the rise and fall of the waters is only
a matter of days. A high flood is very destructive covering the crops and often
entering the homes of the farmers, who are forced to seek shelter on higher
ground. The black mud left behind by the waters is fertile and the ground is
improved by the moisture yet the damage of the flood outweighs its good effects.
Prevention, however, is a difficult matter. The flats are far too extensive and not
Tich enough to pay for the building of levees. Aliso floods are not nearly so
numerous or so high as they used to be. The best method of controlling floods is by
water storage. There are at present three dams on the Nepean which hold back
some of the water at time of flood and, by thus knocking off the peak of the flood
wave, do much good by making the rise of the waters less abrupt. The river is the
one nearest to Sydney and therefore its natural source of water supply, while the
inhospitable character of the rugged mountainous area through which it flows
prevents settlement and keeps the water fairly pure. It is suggested to build a
large dam on the Warragamba River which will assure an ample supply of water
to Sydney and at a time of heavy rain will help to smooth down the flood wave so
considerably that the water may possibly be contained within the river banks.
Dams, however, cannot actually prevent floods and those built on the Nepean have
not been the cause of the decrease in the number of floods experienced at Windsor
of late years. This decrease is rather a factor of declining rainfall for precipitation
on the catchment area has been steadily less ever since the heavy rainfall of 1870.

References.

Davin, T. W. E., and GuTHRIE, F. B., 1904: ‘The flood silts of the Hunter and Hawkesbury
Rivers.” Proc. Roy. Soc. N.S.W., xxxviii, pp. 191-202.

152 FLOODING OF HAWKESBURY RIVER AT’ WINDSOR.

DE BurcH, 1908: “Warragamba River and Water Supply of the County of Cumberland.”
Report to the Water Conservation and Irrigation Commission, Sydney, N.S.W.

HUNTINGTON, E., 1915: “Civilization and Climate.’’ University Press, London.

HUNTINGTON, E., and VisHmr, S. S., 1922: “‘Climatic Changes, their Nature and Causes.’
Yale University Press.

Hunt, 1916: “Results of Rainfall Observations made in N.S.W. during 1909-1914.” Krom
the Commonwealth Bureau of Meteorology.

METROPOLITAN BOARD OF WATER SUPPLY AND SEWERAGE: Thirty-sixth Annual Report for
year ended 20th June, 1924.

Taytor, T. G., 1923a: “The Warped Littoral Round Sydney.’’ Proc. Roy. Soc. N.S.W.,
lvii, pp. 58-79. :

Taytor, T. G., 19238b: “Geography and Australian National Problems.” Report Aus-
tralasian Association Advancement of Science, Wellington, Vol. 16, pp. 4383-487, 1924.

TEBBUTT, 1908: ‘““Windsor Rainfall. A Retrospect for 45 Years.” Windsor and Richmond
Gazette, January, 1908.

WATER CONSERVATION AND IRRIGATION COMMISSION: Annual Report for Year ended June,
1925.

DESCRIPTION OF PLATE VIII.

Figure 1.—Flood silts on a grassy bank at Windsor showing the laminated character
of the deposits.
Figure 2.—Irregular cracks due to drying in the flood silts at Windsor.

NEW GALL-FORMING THYSANOPTERA OF AUSTRALIA.

By Dupitry Movurron.
(Communicated by W. W. Froggatt.)

(Plate ix and fourteen Text-figures. )
[Read 25th May, 1927.]

The gall-making thrips of Australia include some of the most interesting
forms of all the Thysanoptera, and I am greatly indebted to Mr. W. W. Froggatt
who has supplied me with abundant material of many species, several of which
are new. Iam offering this brief paper as a preliminary one and hope it will
help to stimulate other entomologists to collect and study these thrips whenever
they are observed. I wish here to express my appreciation to Dr. H. Priesner for
examining many of these specimens.

KLADOTHRIPS AUGONSAXXOS, n. Sp. (Plate ix, No. 1347.)

This species of Kladothrips can be distinguished from K. rugosus, the one
other known species of this genus, also found by Mr. Froggatt, by the following
characters:

K. rugosus Froggatt. K. augonsaxrxos n. sp.

All antennal segments except the first,
more or less uniformly light yellow to
yellowish brown.

Pterothorax and first two or _ three
abdominal segments yellow, or yellowish
orange. Middle and hind legs uniformly
dark brown.

Third antennal segment, also sometimes
tip of two, yellow, all others dark brown.

Pterothorax and abdominal segments
quite uniformly dark brown. Tips of
middle and hind tibia and tarsi often
yellowish.

The most striking difference is the enlargement of the egg sac in the abdomen
of K. augonsaxxos which I have never before observed among the Thysanoptera.
The abdomen is normal in one specimen before me, apparently before the develop-
ment of the egg sac. Its sides are almost parallel, being reduced gradually from
the sixth to ninth segments and there is no apparent distention of the connecting
tissue between the segments. In other specimens the egg sac is variously enlarged,
although the chitinous walls of the segments do not appear to lose their
original shape. In one specimen which is typical, the connecting tissue between
segments three and four is distended and forms a small, transparent ring
extending entirely around the body. The connecting tissue between segments four
and five is distended into a larger transparent ring which could possibly contain
a single row of eggs, placed end to end. The great spherical egg sac formed
between segments five and six is three or four times the width of the body in
diameter, tough and strong, whitish, almost transparent in colour, and is seen
to contain a great mass of eggs. Segments six to nine and the tube appear to
retain their normal shape and size.

U

154 NEW GALL-FORMING THYSANOPTERA OF AUSTRALIA.

Female holotype—Measurements: Total body length in normal condition
about 2:60 mm., with distended egg sac about 2:75 mm. Head length 0:33 mm.,
width 0-165 mm.; prothorax length 0:33 mm., width 0-36 mm., including prominent
fore coxae, 0°45 mm.; width of pterothorax 0-45 mm.; tube length 0:20 mm.,
width at base 0°08 mm. Antennae, length (width) segment i, 30 microns (39
microns) ; ii, 60 (38); iii, 75 (33); iv, 69 (36); v, 63 (32); vi, 48 (30); vii and viii,
72 (27); total 435 microns. Length (width) of femora, fore femora 416 yu
(133 »), middle femora 135 w (63 «); hind femora 225 uw (90 uw).

Colour dark brown. Extreme tips of fore femora, fore tibia and tarsi
including tarsal claws, also third segment of antenna and extreme tip of two
orange yellow. Fore coxae, pterothorax except outer borders, extreme bases and
tips of middle and hind tibia, and middle and hind tarsi light brown. Connecting
tissue between abdominal segments light brownish yellow, especially conspicuous
after the egg sac begins to form.

Head about as long as prothorax and half as wide, with straight, almost
parallel sides (Text-fig. 1). Postocular spines short and stumpy with dilated tips,

Kladothrips augonsaxxvos, n. sp.
Fig. 1. 9 Head and prothorax, dorsal view.
Fig. 2. @ Right antenna, dorsal view.

situated well back from the eyes. Eyes comparatively small, neither prominent
nor protruding, with small facets, not pilose. Anterior ocellus on apex of head,
posterior ocelli contiguous with inner margins of eyes. Mouth cone _ short,
triangular, reaching hardly one-third the length of the prosternum, with bluntly
rounded tip. Antenna apparently 7-sezgmented. The terminal segment has a
distinct transverse, oblique suture indicating clearly that it is formed by the
fusion of the two terminal segments. (This is also true of the antennae of
K. rugosus Froggatt, specimens of which are before me.) Antennae about one
and one-third times as long as head; segment three club-shaped, segments four,

BY DUDLEY MOULTON. 155

five and six abruptly constricted, each at about one-fourth its length from the
base, these narrowed portions cylindrical, the rest of the segments subglobose.
Segment three with one simple sense cone near tip on ventral side, segments four
and five each with two, and segment seven with one. All sense cones short and
transparent (Text-fig. 2).

Prothorax, not including prominent coxae, somewhat wider than long, shield-
shaped, with a distinct median longitudinal thickening. Prominent spines on
anterior and posterior angles, also on prominent coxae. All spines stout and trans-
parent, with blunt tips, those on posterior angles 75 microns long. Pterothorax
as wide as prothorax and fore coxae combined, sides almost parallel with a
small angular indentation in the middle on either side. Anterior legs much
enlarged, fore femora about one-fourth longer than head and three-fourths as
wide as width of head. Armature of fore tibia reduced to blunt inconspicuous
knobs at end on inner side. Claws of fore tarsi are long, straight and pointed,
as long or longer than length of tarsi, and project at right angles. Middle legs
much reduced, the median femora smaller than the fore tibia; posterior legs
somewhat larger. Wings well developed, entirely clear, not narrowed in the
middle, with ten double fringe hairs along posterior margin near tip.

Abdomen of normal shape before the formation of egg sac. Tube almost
two-thirds as long as head.

This thrips is a true gall maker and produces a spherical gall from one-half
to three-quarters of an inch in diameter, hollow, with an outer shell wall of about
one-sixteenth of an inch in thickness. The gall is a deformity growing out of
the leaf. One female inhabits each gall; she scatters her eggs, hundreds of them,
on the inner surface where the larvae hatch, and apparently feed as I have found
larvae within the parent gall in all stages.

No males have been observed.

Female holotype in author’s collection; paratypes in the Froggatt Collection,
Canberra, Australia. ,

Hostplant: Gallmaker on Acacia doratoxylon.

Habitat: Gilgandra, New South Wales.

CHOLEOTHRIPS, new genus (No. 518).
(Sub-family Kladothripinae. )

Head one and one-third times as long as wide and slightly shorter than
prothorax, broadest across at the eyes, constricted uniformly toward the base.
Cheeks with several thorn-like bristles, a single pair about one-third the length
of the head from posterior margin, long and conspicuous and almost one-half
as long as prominent postoculars. Eyes large. Ocelli present. Mouth cone short,
reaching about two-fifths the length of the prosternum, bluntly rounded, labrum
blunt.

Prothorax shield-shaped. Fore coxae enlarged, prominent, and together
with the prothorax wider than the pterothorax. Fore femora greatly thickened.
Fore tibia stout and armed at the end within with a strong tooth. Each fore
tarsus with two teeth. Wings not constricted sole-shaped, but still a little
smaller in the middle. Tube almost as long as head. Type: Choleothrips
geijerae n. sp.

CHOLEOTHRIPS GEIJERAE, n. sp. (Plate ix, No. 518).

Female holotype-—Measurements: Total body length 2.42 mm. Head length
0-33 mm., width 0:25 mm.; prothorax length 0-36 mm.; width 0-416 mm.; including

156 NEW GALL-FORMING THYSANOPTERA OF AUSTRALIA.

coxae 0:55 mm.; mesothorax width 0-52 mm.; abdomen width 0-416 mm. Antennae:
Segment i 27 microns, segment ii 54 microns, other segments broken off. Legs,
fore femora length 0-41 mm., width 0:20 mm.; middle femora length 0:23 mm.,
width 0:10 mm.; hind femora length 0:33 mm., width 0:12 mm. Tube length 0-315
mm., width near base 0:09 mm.

Colour: Prothorax and fore legs quite uniformly yellowish brown; head, first
antennal segments, pterothorax, abdomen, middle and hind legs dark brown, with
joints and tarsi of middle and hind legs lighter. Wings uniformly smoky coloured.
All prominent spines yellow. (Two paratype specimens have the prothorax and
fore legs dark brown like the rest of the body.)

Head one-third longer than greatest width across at the eyes, constricted
posteriorly (Text-fig. 3). Postocular spines pointed, 90 uw long. Longest cheek

Fig. 5

Fig. 6,

Choleothrips geijerae, n. sp.
Fig. 3. 2 Head and prothorax, dorsal view. Fig. 4. 92 Tip of abdomen, dorsal view.
Fig. 5. 92 Right fore leg, ventral view. Fig. 6. 9? First segment of abdomen, dorsal view.

spines 39 w long. Eyes large, occupying four-fifths the width and about four-
ninths the length of the head. Facets very small, not pilose. Ocelli present,
placed well forward on head, anterior one on apex, posterior ocelli contiguous
with anterior inner margins of eyes; mouth cone short, rounded, labrum blunt.
There are two distinct swellings ventrally, one on either side of the head, just
anterior to the base of the mouth cone. Antennae 8-segmented. Unfortunately the
antennal segments beyond the second joint have been broken from all of the
specimens before me.

BY DUDLEY MOULTON. 157

Prothorax distinctly shield-shaped, one-eighth wider than long but about
one-third wider than long when the fore coxae are included. A prominent
spine on each fore angle, 90 u long and one on each posterior angle 180 uw.
Long spines on angle of fore coxae 135 yw. All spines pointed. The median
dorsal suture terminates in front in a triangular enlargement with distinct
sculpturing. Mesothorax widest, excluding protruding fore coxae of prothorax,
with sides almost parallel. Metathorax with sides tapering gradually to meet
the more slender abdomen. Fore femora greatly enlarged, about four-fifths as
wide as head and nearly one-third longer (Text-fig. 5). Fore tibia enlarged and
armed with a strong curved tooth at the end on the inner margin. Fore tarsi
each armed with two teeth, the larger inner one almost as long as the width of
the tarsus and with a curved tip, the outer smaller one on the ventral or outer
side. The middle and hind legs are much smaller with clearly enlarged femora
but small in comparison with the fore pair. Wings fully developed, reaching
nearly to tip of abdomen with sides almost parallel; twenty-eight accessory
hairs on posterior margin of fore pair.

Abdominal segments three to seven inclusive, are almost equal in length
and width, and have parallel sides. Segment two is subequal in width but
longer. The tergum of segment one is broad in the centre but compressed to
curved points on either side, each bearing a long spine 0:93 mm., with distinet
sculpturing (Text-fig. 6). The tube is almost as long as the head with straight
sides tapering only slightly toward the tip. The long spines of the ninth
segment reach approximately four-fifths the length of the tube, spines on the
end of the tube are somewhat shorter (Text-fig. 4).

Described from twelve females, including holotype, with prothorax and fore
legs distinctly yellowish, and two paratypes with prothorax and fore legs con-
colorous with the rest of the body. Holotype in author’s collection. Several
paratypes in Froggatt Collection, Canberra, Australia.

Hostplant: Geijera parviflora (Wilga), forming rolled leaf galls.
Habitat: Gunnedah, New South Wales (W. W. Froggatt).

DOLEROTHRIPS (?) GEIJERAE, n. Sp. (Plate ix, No. 518.)

Female holotype——Measurements: Total body length 2-48 mm. Head length
0:30 mm., width 0:20 mm.; prothorax length 0°30 mm., width 0:30 mm., and 0°36
mm., including fore coxae; mesothorax width 0°55 mm. Antennae: Segment i,
36 microns, segment ii 54 microns, other segments broken off. Legs, fore femora
length 0-31 mm., greatest width 0-12 mm.; middle femora length 0:21 mm.;
greatest width 0:09 mm.; hind femora length 0:25 mm., greatest width 0-12 mm.
Tube length 0°33 mm., width 0:10 mm. across swollen part near base and 0:06 mm.
just before tip.

Colour quite uniformly deep brown, wings transparent to slightly smoky.

Head almost one-half longer than width across at eyes. Cheeks swollen in
the middle and clearly constricted toward the base, with several short sharp spines.
Postocular spines not conspicuous, set back from eyes about one-third the distance
between posterior margin of eyes and posterior margin of head. Eyes large
occupying two-thirds the width and one-third the length of the head. Outer
border of eyes clear light yellow, facets very small; not pilose. Ocelli present,
placed well forward on head, anterior one on apex but not protruding over basal
segments of antennae, posterior ocelli contiguous with inner anterior margins of

158 NEW GALL-FORMING THYSANOPTERA OF AUSTRALIA.

eyes (Text-fig. 7). Mouth cone short, rounded labrum _ blunt. Antennae
8-segmented.

Prothorax shield-shaped, as wide as long but about one-fifth wider with
the prominent fore coxa included. Spines on fore angles weak, spines on
posterior angles pointed, 60 m. long, several short stout spines on angles of
fore coxa, longest 40 m. Median dorsal thickening fades in front a short
distance before the anterior margin but extends almost to the posterior margin of
the prothorax. Mesothorax with rounded anterior angles, sides of both mesa-
and meta-thorax constricted gradually toward the posterior margins. Fore
femora enlarged, as long as head. Fore tibia unarmed; fore tarsi each with two

[ S™

Fig./] &

Dolerothrips (7?) geijerae, n. sp.

Fig. 7. @ Head and prothorax, dorsal view. Fig. 8. 9? Tip of abdomen, dorsal
view. Fig. 9. @ First segment of abdomen, dorsal view. Fig. 10. @ Tip of right
fore tibia and tarsus, ventral view. Fig. 11. co Tip of right fore tibia and tarsus,
ventral view.

teeth, the larger more conspicuous one extends outward at right angles and is
nearly straight, the terminal smaller ventral one is distinctly curved (Text-fig. 10).
The middle and hind legs are strong, with femora enlarged, but small in com-
parison with the fore femora. Wings fully developed, reaching to middle of
seventh segment, sides almost parallel, 21 double fringe hairs on posterior
margin of fore wing.

Abdomen elongate, first segment broad in the centre and narrowed to a
point on either side, it displays distinct sculpturing and bears a single long
yellow spine at each outer angle (Text-fig. 9); segments two to six increase in
width gradually, and seventh, eighth and ninth with sides rounded and clearly

BY DUDLEY MOULTON. 159

narrowed posteriorly; tergites of segments two to nine more or less distinctly
reticulated; tube slightly longer than head, swollen at the base, narrowed
gradually to a rounded tip (Text-fig. 8).

Male: The male is somewhat smaller, about 1:88 mm. in length, similar in
colour and shape. The fore tarsus is smaller and tarsal tooth is about one-half
as large as in female (Text-fig. 11).

Described from nine females and three males.

Holotype in author’s collection. .Paratypes in the Froggatt Collection,
Canberra, Australia.

Hostplant: Geijera parviflora (Wilga), found in company with Choleothrips
geijerae forming rolled leaf galls.

Habitat: Gunnedah, New South Wales.

EOTHRIPS BURSARTAE, n. sp. (No. 516.)

Female holotype.—Measurements: Total body length 1:66 mm. Head length
0-21 mm., width 0-20 mm.; prothorax length 0-166 mm., width 0°33 mm.; meso-
thorax width 0:38 mm. Tube length 0:183 mm., width 0:08 mm. Antenna length
(width) segment i, 30 microns (33 microns); ii, 48 (36); iii, 54 (36); iv, 48 (39);
v, 48 (36); vi, 42 (33); vii, 48 (27); viii, 24 (18); total length 345 microns.

Hothrips bursariae, n. sp.
Fig. 12. 9 Head and prothorax, dorsal view. Fig. 13. 9 Left fore tibia and tarsus,
ventral view. Fig. 14. 92 Right antenna, dorsal view.

Colour brown to dark brown with tips of fore tibia, all tarsi and segments
three to five and sometimes six lighter. Wings uniformly light smoky grey.
Prominent spines yellow to yellowish brown. Body with numerous orange
pigment spots or blotches visible in light coloured specimens.

160 NEW GALL-FORMING THYSANOPTERA OF AUSTRALIA.

Head broadly rounded with cheeks slightly arched, broadest across middle
line, dorsum with numerous confluent cross striations; postocular spines rather
short, stump with dilated tips; cheeks with several short spines. Eyes occupying
three-eights the length of the head, with small facets, indistinctly pilose. Ocelli
placed far forward, posterior ocelli contiguous with anterior inner margins of eyes,
with dark orange red crescents (Text-fig. 12). Mouth cone reaching posterior
third of prosternum, bluntly rounded labium but with pointed labrum. Antennae
8-segmented, approximately one and two-thirds times longer than head, all seg-
ments comparatively stout, fourth segment almost five-sixths as wide as long. Two
sense cones on segments three, four, five and six, one on segment seven, all are
short, blunt and transparent (Text-fig. 14).

Prothorax almost as long as head but noticeably wider, with sides diverging
posteriorly. Spines on anterior angles short, mid-laterals a little longer, those on
posterior angles longest, all, however, are short, stocky and with dilated tips; each
prominent angle of fore coxa also has a similar blunt tipped spine and two or
three short pointed ones. Pterothorax broadest, sides almost parallel, converging
only slightly near posterior line. Legs rather short, fore femora thickened, fore
tibia unarmed, fore tarsi armed each with a spur-like tooth on inner margin and
a smaller horn-shaped tooth near tip on ventral side (Text-fig. 13). Wings fully
developed, broad, with parallel sides, not constricted in the middle, with ten or
twelve double fringe hairs.

Abdomen almost as wide as pterothorax, first seven segments of equal
width, eight and nine conspicuously narrowed. Tube slightly shorter than head.
Two prominent spines on each posterior angle of segments two to nine, the outer
shorter ones (38 microns on segment two) gradually become longer until they
reach a maximum length of 111 microns on segment nine; the inner ones, 60
microns on segment two, increase to a maximum length of 135 microns on segment
nine, both pairs have blunt or dilated tips. Segments two to seven also have
two additional pairs of curved spines on either side, the posterior pair on each
segment much stronger than the anterior pair. The longest spines on the end
of the tube are two-thirds as long as the tube, 135 microns.

The male is very similar to the female, although somewhat smaller, the
abdomen decreases in width gradually from the fourth segment to the tube.

Described from numerous females and males taken from galls on leaves of
Bursaria spinosa (Native blackthorn) at Warrah and Sydney, New South Wales,
Australia, by W. W. Froggatt.

Holotype in author’s collection. Paratypes in the Froggatt Collection,
Canberra, Australia.

EXPLANATION OF PLATE IX.

No. 518. Rolled leaf galls on Geijera parviflora (Wilga) made by Choleothrips geijerac
and Dolerothrips geijerae. Natural size. ;

No. 1347. Galls on Acacia doratoxylon made by Kladothrips augonsaxxos Moulton
Slightly enlarged.

AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON.
Part iv. HasitTat FACTORS AND PLANT RESPONSE.

By Joun McLucnis, M.A., D.Sc., Assistant-Professor of Botany,
the University of Sydney;

AND

ArtHuR H. K. Perrizt, M.Se., Senior Demonstrator in Botany,
the University of Melbourne.

(Twenty Text-figures.)
[Read 25th May, 1927.]

Contents.
1. Introduction.
2. The Habitat Factors.
Climate of the Region.
Humidity and Evaporative Power of the Air.
The Moisture-Content of the Soil.
The Physical Structure of the Soil.
The Hydrogen-Ion Concentration of the Soil Solution.
3. Plant Response.
Leaf Structure of the Chief Constituents of the Ceratopetalum-Doryphora
Association.
Leaf Structure of the Chief Components of the Hucalyptus piperita-
E. haemastoma Association.
Root-systems.
The Ultimate Conception of the Vegetation of Mount Wilson.
5. Summary.

1. Introduction.

The previous studies (Part i, 1924, Brough, McLuckie and Petrie; part ii, 1925,
Petrie; Part iii, 1926, McLuckie and Petrie) of the vegetation of Mount Wilson
have brought to light many interesting problems, the solution of which demands
intensive study. The most outstanding feature of the vegetation of this region
is the great contrast between different associations occurring in close juxta-
position: sclerophyllous Hucalyptus Forest alternating with Sub-tropical Rain
Forest frequently and sharply in a comparatively small area. This condition has
led us to make a more detailed enquiry into the habitat factors which are
probably responsible for these remarkable distributional features, and the results
of the investigation form the subject of the present communication.

Owing to the absence of a properly equipped permanent field laboratory, the
scope of quantitative investigations was somewhat limited. From what has been
done, however, it is possible to deduce, in a general manner, what are the basic
factors controlling the distribution of the plant communities in this region, and
to give an explanation of some of the remarkable features which were recorded
in the previous papers. A number of more detailed observations are also recorded
upon the ecological structure of the vegetation and upon the nature of its responses
to the environment.

Vv

162 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

We have to record our indebtedness to Mr. D. J. Mares, Commonwealth
Divisional Meteorologist, for rainfall and temperature data which have been
utilized in this paper.

2. THE HapitatT FACTORS.

It is evident that in a small region like that under consideration, certain
factors, e.g. climate, will be practically uniform throughout, and, although stamping
their impress on the nature of the vegetation, are not directly the cause of the
diversity of habitats within the region. Neither rainfall nor the general regional
features of climate and season are variable to any marked extent at Mount Wilson,
although it must not be thought that they are without their effect on the
vegetation; but it is in other directions that one has to seek for the explanation of
xerophytic and mesophytic associations existing side by side, and for the occurrence
of Rain-Forest in a region with a rainfall rather below that of the typical habitat
of that association-type. From what has already been written it will be clear
that the edaphic factors, rather than the climatic, are of fundamental importance
in producing the great differences in habitat which result in the presence of these
two types; and it appears that we have here an example of compensating factors;
in a region in which the climax association-type is sclerophyllous Forest, a local
high edaphic favourableness apparently produces the same resultant as would a
much higher rainfall with a poorer type of soil; so that a “post-climax’’, to use
Clements’s term (Clements, 1916), is here possible in the form of mesophytic Forest.
It has also been made evident, however, that it is only in places where exposure is
favourable that this effect is produced, so that exposure becomes a second factor
of outstanding importance. It is therefore the edaphic factors, and the atmos-
pheric humidity, which depends largely on exposure, that have claimed the
greatest consideration in the present enquiry.

Among the general problems of distribution which it was hoped to elucidate
are the following:

(1) The comparatively small amount of invasion of the basalt by types from
the Eucalyptus Forests of the sandstone.

(2) The reason why the Rain-Forest grows equally well in the sandstone
gullies and on the basalt caps. |

(3) The absence from the basalt of certain Rain-Forest components in the
sandstone gullies, such as Callicoma, Todea, Blechnum capense and Histiopteris
incisa (see Part iii).

CLIMATE OF THE REGION.
Temperature.

No regular records have been taken at Mount Wilson, but the accompanying
averages (Table i) of an eight years’ record at Mount Victoria, which is only a
few miles distant and is of approximately the same altitude, will give a sufficiently
accurate conception of the nature of this factor in the region under study.

From these figures it is seen that the extremes are moderate; and although
the summer months are fairly hot, the comparatively low minimum values in
winter, and the occasional occurrence of snow, are undoubtedly of great significance
in controlling the development of the Rain-Forest. We have described it as a
luxuriant forest; yet it has not the tropical luxuriance of the more northern Rain-
Forests. The floristic composition is comparatively limited, and there are only
two dominants; whereas a tropical forest is characterized by great floristic
variation. The structure is not excessively complex. The common features of a

BY J. MCLUCKIE AND A. H. K. PETRIE. ° 163

TABLE i.
Monthly Temperature Averages at Mount Victoria.

|
| Mean Maximum. Mean Minimum.
Month. Degrees Fahrenheit. Degrees Fahrenheit.
January Reacts: Sey pL ieee 74-7 53:6
TRAUEIRY ca ou Go oe | 73:3 54-3
March .. : | 68:6 51-0
April 5) or. oe oe er eee ee 61:7 46-3
May Akush nical onary et CS MECC | 53:8 41:6
INCOME Ee! pre ys 6 a 48-4 | 37-3
July 5 Sia EApruee Es acne 41-6 | 35-2
August Set UN as ie RIO le 51-6 | 37-0
Sepiemalaee o5 boa G0 oc | 58-8 | 39-6
October JSS eee a an A 65-4 | 44-3
INIOWENM NS oo oo po <6 | 71:5 | 48-0
IDYXOEIIN GVO 56 bo oa los 74-9 50:6
|
Average for year .. =. | 62-4 44-9
|

tropical Rain-Forest, e.g. very large leaves, plank buttresses, cauliflory, or epiphylly,
such as are found even in the coastal forests of the same latitude in New South
Wales, are practically absent. While epiphytic mosses and lichens are abundant,
epiphytes of Pteridophytic and Angiospermic affinity are not profuse. Indeed, we

are driven to realize that the luxuriant physiognomy of the Rain-Forest at Mount
Wilson is imparted not by tropical structure and adaptation, but by its wealth
of Pteridophytes and by its closeness of structure arising from high edaphic
favourableness. Features such as these are in accordance with what one would
expect in a montane sub-tropical or temperate Rain-Forest; and although the
moderate rainfall is an important factor in causing them, there is no doubt that
low temperatures in winter are also effective to a considerable degree.

Rainfall.

While the Blue Mountains as a whole have an average annual rainfall of 30
to 40 inches, Mount Wilson is exceptional in having the higher value of 46 inches.
In Text-fig. 1 is given the monthly variation at Mount Wilson and also that at
Mount Victoria, a more typical Blue Mountain area. It will be seen that not only
is the rainfall at Mount Wilson higher than that at Mount Victoria throughout
the whole year, but during the summer it is much higher: the maximum
precipitation takes place during the hottest months, December to March.

This state of affairs is of considerable importance in connection with the
occurrence of the Rain-Forest at Mount Wilson. While the basalt soil is the main
factor leading to the development of this community on the plateau, there is no
doubt that were it not for the high summer precipitation it would be a much less
favourable habitat for the Malayan Rain-Forest types which at present characterize
it. During the winter when desiccation is less, the smaller rainfall is not
deleterious.

At Mount Victoria, although the maximum rainfall is also in the summer, it
is not by any means so high; and this fact probably explains in part the

164 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

MAR. | APR. | MAY. |JUNE.|JULY.] AUS. | SEP | OCT.] NOV, [DEC | JAN. |

be v
A pyanlt |e

Text-figure 1. Annual rainfall graphs for Mount Wilson and Mount Victoria.

occurrence of Rain-Forest in shallow sandstone gullies at Mount Wilson which at
Mount Victoria would probably be occupied merely by sclerophyllous Hucalyptus
Forest.

The basaltic soil and the comparatively high summer rainfall then, are respon-
sible for the maintenance of the luxuriant vegetation of the Mount Wilson area.

HUMIDITY AND EVAPORATIVE POWER OF THE AIR.
Methods.

At the beginning of September, 1924, measurements were made of the
evaporative power of the air in different habitats in order to obtain some
knowledge of the requirements of the vegetation with regard to this factor.

As standard Livingston atmometer cups were unobtainable at the time,
evyaporimeters were made from Chamberland filter-candles attached to burettes by
a piece of rubber-tubing, the whole being carefully filled with water so as to avoid
the inclusion of air-bubbles. As it was desired to compare daily with nightly
readings in the Rain-Forest to ascertain whether conditions bordering on satura-
tion obtained there, burettes were used in place of the jars more commonly
associated with the Livingston atmometer. This apparatus gave a rapid and
accurate record of the water evaporated; it was found, however, during preliminary
tests in the laboratory, that in a very humid atmosphere the presence of a column
of water in the burette above the level of the filter candle resulted in a tendency
towards exudation of water through the pores of the porcelain. To avoid this in
all the evaporimeters used, the filter was raised above the level of the water in
the burette by the insertion of a vertical piece of glass tubing between the filter
and the burette. After this was done, and so long as the water in the burette was
not above the level of the filter, trials in the laboratory showed that in an atmos-

BY J. MCLUCKIE AND A. H. K. PETRIE. | 165

phere of constant humidity the rate of evaporation was constant, and was not
measurably affected by the level of water in the burette.

Shreve (1914, p. 43) has pointed out that atmometer readings taken in very
dry and very moist climates are not strictly comparable, owing to a difference in
the character of the evaporating water-film; and this objection would probably
apply to our comparisons of readings taken in the Hucalyptus Forests with those
in the Rain-Forests. The error, however, is not likely to be large enough to
influence the general conclusions to be drawn from the results cbtained.

The evaporimeters, although having approximately the same rate of evapora-
tion, were standardized in the laboratory by comparison with the evaporation from
an open water surface, whereby a factor was obtained for each evaporimeter. By
this means the readings were standardized to the evaporation in cubic centimetres
from an open water surface one square metre in area in which form they are
expressed in the results given in Text-fig. 2.

Readings were taken early in the morning and at sundown for five successive
days, the total evaporation for the twelve hours representing day or night being
calculated therefrom. The total daily and nightly evaporation for the whole period
is given for each station in Text-fig. 2.

Discussion of Results.

This comparative study of the evaporative power of the air in different
habitats gives an interesting confirmation of our observations upon the effect of
the exposure on the distribution of the various communities.

The Rain-Forest, as we have previously stressed, is limited to the most
sheltered habitats; and it is seen that the evaporative power of the air within
it is very much less than in any other habitat. Water vapour appears to arise in
considerable amount from the damp soil, forming a humid layer among the ferns
on the floor of the forest. Owing to the deep shade and the extreme absence of
wind in the forest, the moisture-content at the ground-level is high, and must
border on, and no doubt at times attain saturation during the night. The greatly
increased insolation at high levels in the Forest lowers the relative humidity
and causes a striking difference between the values for the two strata expressed
in the diagram. No such dissimilarity is noticeable in the more open communities,
where there is little difference in the insolation of the various strata.

The Alsophila society of the Hucalyptus goniocalyx-E. Blazxlandi association
has been described as much more tolerant of exposure, avoiding only the full brunt
of the westerly winds on the open summits of the basalt caps, which are occupied
by the Pteridium society in the H. Blaxzlandi consociation (Part ii, p. 149). The
results from stations 4, 5, and 6 show that the Alsophila society can withstand
very high atmospheric dryness. The values in stations 5 and 6 are even higher
than that in station 8 in the H. piperita Forest; and, although it is to be expected
that more exposed westerly regions in the latter community would give higher
values still, it is nevertheless evident that the series of increasingly mesophilous
communities is not distributed in a series of habitats characterized by a corres-
pondingly increasing atmospheric humidity. We are therefore led to suppose
that the distribution of the Alsophila society is governed mainly by soil-moisture,
and that it is very largely indifferent to exposure, except perhaps where it reaches
an extreme on the open summits of the basalt caps.

The low values of the evaporation during the night periods are sUsleningant
since the westerly wind prevalent during the day ceases in the evening.

COLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

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BY J. MCLUCKIE AND A. H. K. PETRIE. — 167

THE MOISTURE-CONTENT OF THE SOIL.

A large number of samples of soil were collected at various times at Mount
Wilson, in each case after a moderate spell of dry weather. These samples were
passed through a 3 mm. sieve and the moisture content was then estimated by
drying at 100° C.

Ceratopetalum-Doryphora association
(basalt).

Ceratopetalum-Doryphora association
(sandstone).

Eucalyptus goniocalyx consociation;
Alsophila stratum-society; Blech-
num discolor fern society.

Eucalyptus goniocalyx consociation;
Ptcridium society.

Eucalyptus piperita consociation.

° 10 20 30 40 50

PERCENTAGE MOISTURE IN FRESH SOIL.
Text-figure 38. Graphical representation of the
soil-moisture ranges of the main associations.

The ranges for the values obtained for the soil-moisture of each association
are given in Text-fig. 3. It must be understood that these ranges might possibly
be found to be somewhat greater if samples were taken continuously throughout
the year; they are nevertheless indicative of the striking features of difference
between the habitats of the various communities.

From an examination of Text-fig. 3 it is seen how the range of soil moisture-
content for the Rain-Forests, whether on sandstone or basalt soil, does not vary
beyond 13%. Thus, although a lower minimum may perhaps sometimes be reached,
comparison with the ranges of the other communities at once suggests that the
distribution of the Ceratopetalum-Doryphora association is determined by water-
content of the soil: the association as a whole is apparently independent of soil-
type, so long as it finds the necessary high moisture-content. This view is
supported by the occurrence of the Rain-Forest on other soils, e.g. shales, in other
parts of the State.

The Alsophila society of the Eucalyptus goniocalyx consociation is not
limited to the same extent by moisture-content, the values ranging from 45-2%
near the Rain-Forest to 23-5% at the outer boundary; it is probable, however, that
there is a minimum value for the lower extremity of the range (in the neigh-
bourhood of 20%) for the habitat of this community, beyond which it is replaced
by the Pteridium society. It may be asked why the high value of 45-2% is
attained in this open community: the answer appears to be that the dense fern
stratum may protect the soil from evaporation just as much as the Rain-Forest
tree canopy. If then the soil is capable of holding this high percentage of moisture
it would be expected that the Ceratopetalum-Doryphora Forest would be occupying
the habitat; but, as we have seen, the exposure and the prevailing atmospheric
humidity are not suited to the requirements of the Rain-Forest here, and in a
strong westerly wind this high moisture-content no doubt decreases rapidly.

The Pteridium society appears to be one step further in the succession of the
mesophilous to xerophilous communities; it is also noticeable that the Hucalyptus
goniocalyx consociation is evidently confined to soils of a minimum soil-moisture

168 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

range, which, although low, is much higher than that of the #. piperita
consociation.

The HE. piperita Forest has the lowest values, and there is no doubt, more-
over, that after periods of drought the minimum value would be even lower than
that given. It is thus clear that the sandstone plateau is a highly xerophytic
habitat. It is probable nevertheless that, since water drains rapidly downwards
to the lower levels in sandstone soil, it does not have time to dry out between
falls of rain, more especially as we have seen that the rainfall is fairly evenly
distributed throughout the year.

The comparison of the soil-moisture values of this series of communities
indicates clearly how the zonation of vegetation tends to coincide with a zonation
of soil-moisture. There is no doubt that the falling gradient is increased by the
progressive decrease in the capacity of the vegetation to conserve soil-moisture by
shade and humus production. It appears then that the Ceratopetalum-Doryphora,
Hucalyptus goniocalyz-H. Blaxlandi, and EH. piperita-E. haemastoma var. micrantha
associations are adapted to habitats of decreasing soil-moisture content: each will
occupy the habitat where it finds its range, provided the other controlling factor of
exposure permits; if this does not, one of the following associations in the series
will occupy the habitat. The more xerophytic communities are no doubt excluded
from the more mesophytic habitats by competition.

THE PHYSICAL STRUCTURE OF THE SOIL.

Samples of soil were taken at a depth of six inches from a number of typical
habitats, and on arrival at the laboratory they were passed through a 3 mm. sieve.
Two portions were then removed, the one being dried at 100° C. and ignited for
the estimation of humus, the other being analysed mechanically by the method
adopted by the British Agricultural Education Association (see appendix to
Russell, 1921). The smaller separations, such as coarse and fine sand, were not
made, as only the salient features of difference between the various soils were
required. The results of these determinations are given in Table ii.

Discussion of Results.

Obviously the great difference in the physical structure between the basalt
and sandstone soils will be a key to a number of other factors, since it will affect
the aeration, moisture-retaining capacity, colloidal properties, etc., and will also
be likely to have a marked influence on the root-systems. Aeration is one of
the chief concomitants of the physical structure of a soil; and it is well known
that sandstone, open and porous in nature, is abundantly ventilated and well
drained, while clays, on account of the smallness of the component particles are
poorly aerated and poorly drained, and tend to be infertile through lack of calcium
carbonate. <A priori it might be expected that the basalt, being a clay soil,
although not deficient in calcium carbonate, might have an unfavourable influence
on plant life on account of lack of aeration. Humus in amounts of 10% to 30%,
however, as is present in the upper layers of the basalt soil (samples 1 to 8)
counteracts the baneful effects of clay so far as low aeration goes, and renders
the texture more open, without depreciating its moisture-retaining properties
(Russell, 1921). This consideration probably explains the presence of the same
association on the basalt slopes and in the sandstone gullies throughout this area
—a distributional phenomenon which might lead one to deny the influence of
edaphism on the distribution of vegetation. From this aspect of the huimus-

No. of
Sample.

14

15

Results of Mechanical Analysis of Soil Samples.

Community.

BY J. MCLUCKIE AND A. H. K. PETRIE.

TABLE ii.

ee)

Ceratopetalum-
Doryphora
association.

Hucalyptus
goniocalyx
consociation ;
Alsophila society. |

Do. Tree-fern
boundary.

Do. Pteridium
society.

FE. piperita
consociation.
Pteridium society.

EH. piperita
consociation.

Ceratopetalum-
Doryphora
association.

169

Fine
Habitat. Gravel. Sand. Silt. Clay Humus
|
; PT: | GE TRE, Yo ie ae
3D 21-06 2°34 60-32 13:77
| = = = = 19-04
| -Basalt. — | = — = 26-45
| — | — —- — 19-04
Do. 1 ft. deep. es ees —— = 10-40
Do. 5 ft. deep. wale a ut — 12-11
| Basalt. lees ae = — 30-25
|
Edge of basalt. ares | ales = bea 16-48
Tides joey Ne: Sloped Ni g-e70m |p 68:15 9-97 2:97 | 10-24
below basalt. j
eee | at — 10-04
Sandstone; centre
of plateau. 10-39 59-32 8:45 2-74 19-10
5:28 64:90 2-45 7-75 19-63
Sandstone. 6:80 75:19 6°33 0:00 12-84
Westerly slope re- 13-54 73:40 3:27 1-24 8-55
corded in Part iii,
p. 108, where
Telopea appears.
Same slope; sand- Ba) |) ees 6-11 2-77 5-27
stone below
basalt; floristic
composition,
Part iii, p. 108. |
aes of sandstone $22 Ce Hee ae ot
lateau. rTS¥: TE “e Re:
ive 13-73 60-65 6-26 0-00 19-88
| senasone 1-80 70:58 2-66 9-32 15-64
Sandstone a _ little 9-78 54°35 5:75 2°73 27-39
below basalt.
Except where otherwise stated, samples were taken from a depth of six inches.
Diameter of Particles in various Fractions. ee
i ZEVC Ars
Tine gravel = above 1 mm. PAARL AL ™
Sand = 1-0-037 mm. PNT ee ee of
Silt = 0:037-0-002 mm. Pa eee)
Clay = below 0:002 mm, — fy
poy >
! |
i 7

170 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

content, which appears to counteract the effect of other components, the soils are

perhaps not so fundamentally different in their influence on vegetation. Conse-
quently it is not suprising that the actual number of plants with soil preferences
were found to be very few (Part iii). Such preferences as appear to exist are
probably in most cases the results of differences in properties of the soil solution.

Although the humus will have this striking effect near the surface, at depths
of some feet its content in soils decreases rapidly, as is indicated in the present
instance in samples 5 and 6, and eventually becomes quite low. At the same time
the basalt soil is deep, since the rainfall is such as to cause rapid and penetrative
weathering, and water and organic acids from the humus percolate through the
soil and decompose the underlying rock. It is clear then that at greater depths
the soil will not receive the beneficial effects of a high humus content; so that
the vegetation would be expected to root in the upper regions if it is a type
adapted to a well aerated soil. This point will arise again for discussion in a
later portion of the paper.

The large percentage of clay and humus in sample 1 is plainly the key
to the high moisture-retaining capacity, and consequent mesophilous vegetation,
of the basalt soil. Probably, however, in the surface layers the humus is more
important in this respect than the clay; for, as is usually the case, the moisture-
content closely follows the humus-content. Text-fig. 4 shows that the ranges of

I
Ceratopetalum-Doryphora association H
ae CREO |
t
Ceratopetalum-Doryphora association igtiiiet eres
(sandstone).
Eucalyptus goniocalyx consociation;
Alsophila stratum-society; 3lech- riggs dale
num discolor fern society. |
Eucalyptus goniocalyx consociation;
Pteridium society.
Eucalyptus piperita consociationu,
Ee
i
re) 10 20 30 40

PERCENTAGE OF HUMUS.
Text-figure 4. Graphical representation of
the humus-content-ranges of the soils of

the main associations.

these two properties correspond in a striking manner. It is thus the water-
retaining capacity of the humus which makes it possible for the soil-moisture of
the sandstone habitats clothed by Rain-Forest to be as high as those of the basalt,
despite the absence of clay from the sandstone soil. It must be added, however,
that the stations in the sandstone Rain-Forest from which samples were taken
for moisture estimations were more particularly in the erosion channels occupied
by Dicksonia and Todea, no samples unfortunately being taken from the apparently
drier slopes devoid of tree-ferns; and, moreover, the Rain-Forest on the sandstone
occurs only in valleys, where water is continually draining down from the high-
lands. In this connection it is interesting to note that, as was mentioned
previously, the Rain-Forest at Mount Wilson occurs in gullies which on the other
parts of the Blue Mountains would probably be occupied by a much more
xerophytic community, since the water supply would not be sufficiently constant.
In such habitats the gravitational water drains away rapidly and during a dry
period disappears from the heights and so causes the gullies to become dried

BY J. MCLUCKIE AND A. H. K. PETRIE. Abra

up. Hence the suggestion is offered that the water supply for the gullies at
Mount Wilson is to some extent kept uniform and continuous by slow drainage
from the basalt caps; owing to the low capillary power of such heavy clay soils
the water would be conserved after rain and would be allowed to drain away
slowly and uniformly, thus maintaining a constant supply for the gullies below.

This does not depreciate the previous statement that the humus is the main
factor controlling the moisture-content of the soil, since these two factors are
reciprocal: the formation of a water-retaining Humus depends on the nature of
the aspect and the vegetation allowing an accumulation of moisture in the soil:
and were it not for this humus the water would probably dry out from the
sandstone soil. The relationship between moisture-content and humus is probably
responsible also for differences in the properties of the organic portion of the various
soils. The moist volcanic soil and the moist and well-aerated and well-drained
soil of the sandstone gullies must favour the production of a “mild humus”, as in
such habitats earthworms, fungi, and bacteria flourish; in particular the effect of
earthworms is important since large species are abundant on the basalt and
prove an important factor in ventilating the soil. On the other hand the dry
sandstone soil forms a very different medium for the existence of these organisms,
so that little humification goes on; in this case, therefore, although 7% to 8% of
organic matter was found it is improbable that it would have the same moisture-
retaining capacity as that of the more humid habitats, and therefore is contributing
to the xerophytic nature of the sandstone soil. This absence of humification in
the habitat of the Hucalyptus piperita-E. haemastoma var. micrantha association
adds point to its facies, as the bare patches between the individual plants are
usually occupied by fallen leaves, twigs and branches, which in a more humid
forest would soon rot away, but which here dry and accumulate until they provide
material for the next forest fire.

A consideration of the analytical results shows that apart from the basalt
there is no fundamental difference in the physical structure of the other soils.
There is no evidence of mixing of the two main types except in the case of the
basalt which contains a fairly high extraneous sand fraction, probably blown
in by winds. The clay and silt fractions of the sandstone soils are largely
composed of ferric oxide, a common mineral in the Hawkesbury sandstone, and
which comes down with these fractions in the analysis. Samples such as 9 and 20
show that even on steep slopes there is no evidence of basalt soil being carried
down very far and mixed with the sandstone, although there is no doubt that a
mixture occurs in the ecotone region of the Rain-Forest referred to in Part iii
(p. 98); the main erosion of the basalt, however, takes place in definite drainage
channels, and has been described already (Part iii, p. 105).

In Part iii (p. 108) a description was given of the extension of the Hucalyptus
piperita consociation to the basalt on a westerly slope. This spread is apparently
only a small one, since the analysis of sample 14 shows that the Lomatia society is
actually on the sandstone side of the junction, although from field observations we
concluded that the soil was basalt.

The salient feature of this part of the investigation is the striking corres-
pondence between the ranges of the moisture- and humus-contents of the soil.
The clay fraction of the basalt soil holds sufficient moisture to provide a suitable
substratum for the development of a rich humus, and is aided by shade and
humidity in the Rain-Forest. This explains the mesophytic structure of the
vegetation of the basalt soil, a vegetation which extends on to the sandstone only

172 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

on slopes or in valley floors which are damp through drainage. The predominant 5;
effect of the basalt soil on the distribution of the various associations depends
almost solely upon its capacity for retaining water.

Up to this stage then we seem to have found two outstanding factors
controlling the distribution of the plant associations. The first of these is
aspect, which has both a direct effect on the vegetation through the atmospheric
humidity, and also an indirect effect by influencing the evaporation of water from
the soil. The second factor is the moisture-content of the soil, dependent on the
humus, which in turn is dependent on the clay fraction of the basalt soil, upon
drainage and upon shade.

THE HYDROGEN-ION CONCENTRATION OF THE SOIL SOLUTION.

On account of the high base-content of basalt soils, and the continuous filtering
of water through it, it was thought probable that leaching would have an influence
on the soil acidity; in any case the latter was likely to be a factor of interest in a
region so marked as Mount Wilson in edaphic variation.

Clark and Lubbs’ indicators were used for the estimations of the hydrogen-
ion concentration of the soil. The procedure was based on Wherry’s field-method
(Wherry, 1920), with certain modifications: thus it was found necessary in the
case of soils which remained turbid for a long period to filter them through ashless
filter-papers, having carefully leached these until the washings gave a neutral
reaction with brom-thymol blue; it was also found advisable to use the same
quantity of water for each extract in order to obtain comparative results, a
conclusion arrived at also by other workers (Wherry, 1924).

In Table iii are given the P, values for samples taken at a depth of 6 inches
from the various habitats. Each represents the results of a number of deter-
minations from different parts of the community. The acidity was found to be
practically uniform in these habitats at depths from 2 to 8 inches.

TABLE iii.
Hydrogen-Ion Concentration of the Soil Solution from the Habitats of the
Main Communities.

me
Ceratopetalum-Doryphora association; basalt .. .. doa) PE 7/O10)
Eucalyptus Blaxlandi consociation; summit of basalt cap. oa, Web)
E. goniocalyx consociation; Alsophila society .. .. .. .. 6:5
E. goniocalyx consociation; Pteridiwm society BOS Tea rel. ORS eb)
EH. piperita consociation we Pchiec MMLUCCGT NERS AALL | Asay a ent” cud)
Ceratopetalum-Doryphora arneseniten 2 sandstone ad) bay gree] Pee ROLD

The lack of variation in these results is a prominent feature. Despite the high
content of bases, such as calcium, characteristic of basalt soil, the buffer action
apparently prevents any fluctuation in the hydrogen-ion concentration of the soil
solution. In all the soils except the dry sandstone the high content of colloidal
material in the form of humus and also of clay in the case of the basalt, readily
explains this buffer action; in the case of the sandstone it is to be presumed that
the absence of an appreciable quantity of basic material on the one hand, and the
absence of much humification or abundance of respiring root-systems on the other,
mitigate any tendencies towards departure from neutrality in the soil-moisture.

From this it is clear that the hydrogen-ion concentration of the soil solution
is not a factor of significance in the study of the distribution of the vegetation
at Mount Wilson.

BY J. MCLUCKIE AND A. H. K. PETRIE. 173

3. PLANT RESPONSE.

In preceding communications on the ecology of the Mount Wilson area,
we have described the main features of the physiognomy and distribution of the
flora and have endeavoured to interpret these as responses to certain factors or
factor complexes of the environment. In the present paper we have endeavoured
to analyse the environmental factors and to deduce the relative significance to
the flora of each of the more important. We have indicated the outstanding
differences in the various plant associations which occur throughout this area,
and now furnish an account of the chief anatomical characters of the most
important types of these associations. As Cannon (1924, p. 103) points out, two
different and important groups of factors must be considered in the attempt to
interpret the anatomical peculiarities of plants from the physiological point of view,
namely heredity and immediate environment. The fact that different genera
of the same family and even different species of the same genus attain to harmony
with the environment by the most varied anatomical characters, seems to us
to indicate that another set of influences of an hereditary nature is acting
simultaneously with the present environment, and that the structure of the
living types is, generally speaking, the result of the impress of two different
categories of forces. On the other hand the physiologist knows on experimental
grounds that certain characters are definitely plastic and modifiable by changing
environment; hairiness may be replaced by glabrousness by cultivation in a more
humid atmosphere; sclerophylly may be replaced by more mesophytic characters
under similar conditions; nanism may give way to a taller habit; in short most
of the characters of a highly xerophytic plant are changeable under a changed
environment, provided it acts for a sufficiently long period during the develop-
ment of the individual. The different “adaptations” of plants to the more
xerophytic or mesophytic conditions appear to us to support the view that the
present structure is the resultant of factors now operating but modified in certain
respects by the cumulative influence of hereditary forces. This is all the more
reasonable when we consider that members of the same family growing under
precisely the same conditions in the same locality, show, amidst a general similarity
of response certain individual differences. Speaking generally the types from the
Ceratopetalum-Doryphora Rain-Forest (as the following descriptions will show)
have thin unwettable cuticles, little or no sclerenchyma, stomata flush with the
epidermis and without the outer vestibule characteristic of xerophytic types, poorly
developed palisade and a great development of spongy mesophyll with large air
spaces. The types from the Eucalyptus piperita-H. haemastoma association on the
dry exposed sandstone habitat have stomata sunken in grooves or pits, hairiness,
considerable sclerenchyma as hypoderma or as girders associated with the veins,
strongly developed palisade and proportionately poorly developed spongy mesophyll,
tannin deposits, enormously thick cuticles and outer epidermal walls, and
frequently oil sacs.

In these xerophytic types the outer epidermal wall is generally heavily
thickened owing to the deposition of anhydrides under the influence of the
desiccating conditions. Strong cuticularization is a prevailing feature as the
deposition of cutin is intimately associated with the transpiratory activity of
the leaf. In the species of Banksia mentioned the stomata occur in pits, the
cuticle of which is poorly developed. The stomata are usually sunk below the
general surface of the leaf, and the cuticle forms, in many types, a prominent
ridge overarching the guard cells, thus constricting the approach to them.

174 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

Between these two extreme groups of structures are those displayed by
components of the Eucalyptus goniocalyx-E. Blaxlandi association which develops
on a much less xerophytic habitat than the HL. piperita-E. haemastoma association.
The anatomical characters common to these components of the associations at
Mount Wilson may be seen more readily from the following descriptions and

figures.

LEAF STRUCTURE OF THE CHIEF CONSTITUENTS OF THE Ceratopetalum-Doryphora
ASSOCIATION.

Fieldia australis. The chief characteristics are (a) the very thin cuticle for
facilitating cuticular transpiration, (0b) the poor development of palisade tissue
owing to growth in a shaded environment, (c) the extensive intercellular space
system of the leaf facilitating aeration and transpiration (the air channels being
internal evaporating surfaces), (d) the poorly developed vascular tissue, (e) the
numerous short hydathodes for secretion of excess water, and (f) the numerous
stomata on the lower surface for aeration and passage of water vapour. These
are structural features which characterize a type growing in a shaded moist
environment, where aeration and transpiration require to be stimulated as far as
possible by anatomical deviations from the normal type (Text-fig. 5, 5a). It has
been shown earlier in the paper that in the lower strata of the Rain-Forest, which
form the habitat of Fieldia, the atmospheric humidity is often in the neighbour-
hood of saturation.

Text-figure 5. Section of leaf of Fieldia australis. x 115.
Text-figure 5a. Hydathode of Pieldia australis. x 300.
Text-figure 6. Section of leaf of Atherosperma moschatuwm. x 115.

Atherosperma moschatum. In this type the cuticle and outer epidermal cells
are thin; the stomata are numerous and on the lower surface only. A hypodermal
layer of relatively large clear cells is present on the upper surface, suggesting
an aqueous tissue in structure; a narrow palisade layer is developed, while a
loose spongy mesophyll occupies more than half the cross section of the leaf.

BY J. MCLUCKIE AND A. H. K. PETRIE. 175

The palisade layer contains relatively more chromatophores than the cells of the
spongy mesophyll (Text-fig. 6).

Doryphora sassafras also has a hypodermal layer of clear thin-walled cells
devoid of chlorophyll and suggesting an aqueous tissue. This may be correlated
with the fact that Doryphora frequently grows in more illuminated and drier
situations than prevail in true Rain-Forest areas. The palisade layer is narrow
with considerable spaces between certain cells, and large lysigenous oil-glands.
The spongy mesophyll occupies more than half of the cross section of the leaf
as in Atherosperma. The stomata are numerous and occur on the lower surface.
The cuticle on both surfaces is thin (Text-fig. 7).

Text-figure 7. Section of leaf of Doryphora sassafras. x 115.
Text-figure 8. Section of leaf of Ceratopetalum apetalum. 5 Lala,

Ceratopetalum apetalum. There is no hypodermal layer below the upper
epidermis as in Atherosperma and Doryphora, while the palisade layer is slightly
more elongated in comparison with those types. The spongy tissue is extensively
developed, while a few isolated tannin cells are distributed through it. A few
comparatively thick-walled cells surround the vascular bundles. The cuticle
on the upper surface is more strongly developed than in the previous types
(Text-fig. 8).

Quintinia Sieberi. This is also a typical mesophyte which frequently starts
development from seeds deposited on the trunks of treeferns etc. Compared

OS a oe
eareherare

bpe@ae aT

Text-figure 9. Section of leaf of Quintinia Sieberi. x 115.
Text-figure 10. Section of leaf of Callicoma serratifolia. x 115.

176 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

with Atherosperma, Doryphora, and Fieldia, the palisade is more strongly
developed, although the proportion of palisade to spongy mesophyll is only very
slightly different. Thick walled cells are absent from the smaller veins, while
stomata are both numerous and flush with the other epidermal cells (Text-fig. 9).

Callicoma serratifolia occurs in water courses and drainage channels through-
out the sandstone and consequently shows an anatomical structure intermediate
between the mesophytes of the Rain-Forest and the xerophytes of the Hucalyptus
piperita forest. The upper cuticle is much thicker than in Ceratopetalum. 'The
hypodermal layer contains a tannin-like deposit, the palisade layer is better
developed and occupies more than half of the cross section of the leaf. Air
channels are extensive, stomata are numérous, and hairs are developed from the
lower epidermis. Numerous thick-walled cells occur on the adaxial and abaxial
surfaces of the vascular bundles. These are characteristics which stamp Callicoma
as a form somewhat intermediate between the Rain-Forest Flora and the highly
xerophytic individuals of the exposed sandstone habitat. (Text-fig. 10).

LEAF STRUCTURE OF THE CHIEF COMPONENTS OF THE Hucalyptus piperita-
E. haemastoma ASSOCIATION.
Telopea speciosissima. The upper cuticle is very thick and laminated, the

outer epidermal wall is also thick, and the palisade tissue is composed of
elongated cells occupying about half the cross section of the leaf; stereids occur

Text-figure 11. Section of leaf of Telopea speciosissima. x 115.
Text-figure 12. Section of leaf of Persoonia salicina. x 115.

as strengthening cells both in the palisade and spongy mesophyll. The latter is
composed of small cells in the centre and more elongated cells within the lower
epidermis somewhat suggesting a palisade layer. The cuticle of the lower
epidermis is very papillate and forms a cuticular ridge over the stomata which
are sunk slightly beneath the other epidermal cells. A small outer and large
inner vestibule occur in association with the guard cells—a character to be
expected in a xerophytic sandstone type (Text-fig. 11).

BY J. MCLUCKIE AND A. H. K. PETRIE. ° 177

Persoonia salicina is an isobilateral type in which palisade tissue is developed
on both surfaces, with larger air channels than might be expected in a xerophytic
type. The leaf is thick and slightly succulent, and intensely green owing to the
enormous chlorophyll content of the cells. The centre of the leaf is occupied by
large colourless cells with mucilaginous sap, and cells with thick stratified
mucilaginous walls. Isolated stone cells also occur. The guard cells are not so
sunk as in Telopea, possibly owing to the vertical position assumed by the lamina
through the twisting of the petiole through an angle of 90°. The cuticle is not
nearly so thick as in Jelopea, and forms a smooth layer on the surface (Text-
iiez, 1074) .

Banksia serrata is one of the most extreme xerophytic types in this associa-
tion. The upper epidermis is covered by a very thick smooth cuticle; a hypo-
dermis of two or three layers of clear cells probably represents an aqueous tissue.
The palisade tissue is developed in relation to the cavities on the lower surface,
and is composed of long narrow cells with large air spaces between them. The
cavities occur at frequent intervals and are lined by epidermal cells, interrupted
by raised stomata, and producing twisted hairs. Directly beneath the cavities
there is developed a very loose network of mesophyll cells containing few chloro-
plasts compared with the palisade cells. The vascular bundles are strengthened
by means of sclerenchyma extending as girders from one epidermis to another.
The lower epidermis, excepting that of the cavities, is strongly cuticularized while
a hypodermal layer of colourless cells is situated beneath the epidermis (Text-

fig. 13) (compare Hamilton, A. G., Aust. Ass. Adv. Sci., 1907, Adelaide, p. 484,
1908).

ai

es

stnepet
Soe eek
6 =. sale

re 5
pou
J wi

|

Text-figure 13. Section of leaf of Banksia serrata. x 115.
Text-figure 14. Section of leaf of Persoonia acerosa. x 115.

Persoonia acerosa. The cuticle of this pinoid type is fairly thick, and the
guard cells are depressed at the base of the outer vestibule formed by the cuticular
ridges. The palisade tissue is arranged concentrically within the epidermis. The
centre of the leaf is occupied by the vascular bundle and by colourless cells,
some of which contain starch, and others probably represent aqueous tissue.
Isolated groups of sclerenchyma cells occur in the mesophyll, and in relation to
the vascular bundles. The air space system is more elaborate than might be
expected in a xerophytic type (Text-fig. 14).

Ww

178 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

Banksia spinulosa. In this type the leaf margin is recurved, and numerous
slightly raised stomata occur in the cavity so formed. Hairs arise from the
epidermal cells in the vicinity of the guard cells, which open into fairly large
sub-epidermal chambers or cavities. Very little tissue of a spongy nature is
developed, while about three-fourths of the cross section of the leaf is composed
of elongated palisade cells with very narrow intercellular spaces. The cuticle is
strongly developed on both surfaces, while the outer epidermal wall is thickened;
the hypodermis is composed of deeply staining tannin cells and sclerenchyma.
All the anatomical characters of this type confirm the xerophytic structure of
the leaf to be expected from the habitat of the plant (Text-fig. 15).

Hakea dactyloides is an isobilateral type with strongly developed palisade on
both surfaces. The main bundles are surrounded by sclerenchyma developed as
girders connecting the surfaces of the leaf. The centre of the leaf is occupied
by colourless loosely arranged cells, while the stomata are deeply depressed
and the ridges of the thick cuticle form comparatively large cavities above the
guard cells. Many of the epidermal cells contain a deepiy staining substance like
tannin (Text-figs. 16, 16a) (compare Hamilton, A. G., Proc. Linn. Soc. N.S.W.,
1914, p. 152).

Text-figure 16. Section of leaf of Hakea dactyloides. x 115.
Text-figure 16a. Surface view of cuticular ridges overarching the guard
Céellss xX eLs:

BY J. MCLUCKIE AND A. H. K. PETRIE. ' n9

Eucalyptus piperita has an isobilateral type of leaf, with compact palisade
tissue and narrow air spaces. Stomata are found on both surfaces, while aqueous
tissue occupies the centre of the leaf on each side of the main vascular bundles.
Sclerenchyma is formed in association with the chief vascular bundles and forms
a girder across the leaf. A layer of water-jacket cells is frequently present on
each side of the vascular bundles. The cuticle is very thick, but the outer
epidermal wall is exceptionally thin (Text-fig. 17).

: ——— oo \

Hl Mh) iat
a Hh Me

! a

iA
: mn

He

un a

Sy
ae aii

\
i

Text-figure 17. Section of leaf of Hucalyptus piperita. x 115.
Text-figure 18. Section of leaf of Hucalyptus haemastoma. x 115.

Eucalyptus haemastoma resembles #. piperita in its isobilateral structure.
The chief differences are in the greater development of the cuticle, and the definite
central aqueous zone between the palisade of the lower and upper surface. The
guard cells of this type like those of H. Blaxlandi are distinctly elevated, but
protected by a well-developed outer vestibule formed by the enormously thick
cuticle. The very narrow intercellular spaces and the consequent compactness
of the palisade cells of this species are definite indications of the necessity of
rigid control of the transpiring surface, both internal and external. The large
proportion of oil glands in these two species may be regarded as an indication
of their habitat being more xerophytic than that occupied by the species
H. goniocalyx and H. Blaxlandi in which the glands are less numerous in the
leaf (Text-fig. 18).

The components of the H. piperita-E. haemastoma association have adopted
sclerophylly instead of succulence as their adaptation to the environment. The
majority of the species are glabrous and strongly cutinized epidermal cells take
the place of trichomes. Several of the species are isobilateral, and develop palisade
tissue on both surfaces. The intercellular space system of the mesophyll is
exceedingly variable; but generally speaking, types with strongly developed
palisade tissue have relatively small and narrow intercellular spaces. In ‘he
spongy parenchyma of dorsiventral leaves the spaces are comparatively larger.
Perhaps these features of the photosynthetic tissue may be correlated with the
aridity of the habitat.

180 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

The structure of the foliage of the species of Hucalyptus from the various
associations characterized generally by differences of environment, indicates some
degree of correlation between the anatomical structure and the more important
environmental factors, almost amounting to direct cause and effect.

In #. goniocalyx the palisade tissue is loosely arranged with numerous inter-
vening spaces for aeration, while central aqueous tissue like that in
E. haemastoma is generally absent, except in association with the vascular
bundles. The cuticle and outer epidermal wall are not so extremely developed

aes

NU
\ iy
LG
\ )

Text-figure 19. Section of leaf of Hucalyptus goniocalyx. x 115.
Text-figure 20. Section of leaf of Hucalyptus Blaxlandi. x 115.

as in EH. haemastoma, but nevertheless they indicate a xerophytic type, despite
the fact that it grows in a soil with a high water content. Owing to the height
attained by the individual and the consequent high rate of transpiration due to
exposure and the general evaporating effect of the atmosphere water control is
very necessary (Text-fig. 19).

In Eucalyptus Blazlandi the same structural features are revealed. As in
E. goniocalyx, the guard cells of the stomata are practically flush with the other
epidermal cells, or slightly raised and situated at the base of a cavity formed
by the overarching cuticle (Text-fig. 20). In these two species of Hucalyptus
the stomata are much more numerous than in the sandstone species, and taken
as a whole the anatomical characters of their foliage stamps them as being
types growing under much more favourable conditions of water-supply than
E. piperita and E. haemastoma.

As compared with H. haemastoma and E. piperita the number of oil glands
per unit area is less—a fact which might be interpreted as having some relation
with the difference in the aridity of the habitat.

In E£. haemastoma the colourless aqueous tissue forms a definite zone in the
centre of the leaf between the abaxial and adaxial palisade.

-BY J. MCLUCKIE AND A.

H. K. PETRIE.

181

Table iv gives the relative number of stomata in some of the chief components
of the various associations:

TABLE iv.

Stomata per sq. mm.

Ceratopetalum- E. goniocalyxz- E. piperita-
Plant. Doryphora EF. Blazxlandi EE, haemastoma
Association. Association. Association.
Upper Lower Upper Lower Upper _ Lower
Surface. Surface. |Surface. Surface.| Surface. Surface.
Eucalyptus goniocalyx .. (Isobilateral) 33 109
E. Blaxlandi (Isobilateral) 184 188
Alsophila australis 0 99
Acacia melanoxylon (1sobilateral) 236 246
Lomatia longifolia .. 0 205
Drimys dipetala 0 280
Pittosporum undulatum 0 184
Blechnum discolor .. 0 60
Dicksonia antarctica 0 113
Eucalyptus piperita (Isobilateral) 60 107
EH. haemastoma (Isobilateral) 81 Bi
Persoonia salicina .. : (Isobilateral) 24 28
Telopea speciosissima .. .. 0 184
Hakea dactyloides .. (Isobilateral) 134 140

THE ROOT-SYSTEMS.

The relation of root-reaction and root-type to species distribution is a most
important feature of ecological study, and yet, on account of the difficulty of
investigation, it has been too much neglected. In the present instance, although
no detailed drawings have been made, we are able to place on record some
qualitative observations which have proved of interest.

‘The Rain-Forest of the basalt soil is characterized by the superficial root-
systems of all its components. The root development of the tree-ferns is meagre
and shallow; that of the trees, although spreading broadly and considerably
branched, is confined to a great extent to the first feet of the soil. This is clearly
a reflex of the conditions of aeration as previously described: the lower layers
are probably quite unfavourable for root-growth; at all events no types at Mount
Wilson appear specially adapted to deep penetration in this soil.

In the sandstone Rain-Forests the root-systems of the trees may be some-
what deeper: upon this point our observations are incomplete. It is worthy of
note, however, that Callicoma serrdtifolia, in exception to the other types, has a
deeply penetrating root-system.. This gives some insight into the absence of this
type from the basalt Rain-Forests: if a deep root development is a rigid character,
it is obvious that Callicoma could not grow on the heavy basalt soil without
being suffocated.

Although this suggestion presents itself in the case of Callicoma, it has
nevertheless, still to be explained why TJodea barbara, T. Fraseri, Biechnum
capense, and Histiopteris incisa occur in the Rain-Forest of the sandstone but
not on the basalt; and it must be confessed that no satisfactory interpretation of

182 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

this has been found. The following observations, however, are placed on record
in the hope that they may be of use in subsequently arriving at the solution.

Todea barbara and Histiopteris in the Rain-Forest are mainly confined to
rivulets; such rivulets have not been observed in the basalt Rain-Forests; but one
specimen of Todea barbara was seen in a creek running over basalt soil in a
Eucalyptus-Alsophila community on the south-eastern side of the plateau. Todea
barbara will occur in creeks outside the Rain-Forest in highly isolated habitats
on the sandstone; it has also been found, however, in deep shade under damp
rocks where there was no trace of running water. It may be that the chemical
composition of the soil solution has some influence on the distribution of these
types, but this is a fact which still awaits intensive study.

The Hucalyptus piperita-E. haemastoma var. micrantha association of the
sandstone is characterized by the extremely deep root-systems of most of the
components. It has already been explained that there is probably a higher
moisture-content in the lower strata of the sandstone soil, and it is now
evident that the plants growing thereon penetrate deeply for the apparent purpose
of obtaining this; the open texture of the soil permits them to do so without
suffocation. By this means not only are the roots enabled to obtain sufficient
moisture, but they escape from what is likely to be a high soil temperature in the
upper strata during the summer season. This character is a common feature of
sclerophyllous plants, as has been shown by Cannon (1924). The absence of
these sandstone plants from the basalt, as in the case of Callicoma, is perhaps
due to the character of the root-system: it may be that a deep root-system is a
rigid character here, which, as Cannon (1915) has suggested, is often the case.

Observations on the components of the Hucalyptus goniocalyz-H. Blaxlandi
association have not been made, but it has been found that H. Blaxlandi growing
on basalt soil has a shallow root-system, a fact perhaps correlated with its
preclusion from the dry sandstone.

On the whole, then, it appears that the character of the root-system is a
significant factor in the distribution of the vegetation of Mount Wilson. In the
physiological reaction to their environment the roots of the sclerophyllous sand-
stone types have had to penetrate deeply and this character seems to have
eventually become rigid: the Rain-Forest types have found it better to exploit
the superficial strata where they could obtain all the moisture and nutritive
substances required. The different plants appear to be confined to those habitats
to which their particular type of root-system is adapted.

4, THE ULTIMATE CONCEPTION OF THE VEGETATION OF MouNT WILSON.

Few montane forest regions offer such a unique field as Mount Wilson for
ecological study, provided as it is with two such widely different floras and with
two such widely different habitats. The main basis for our study has, indeed,
centred round the way in which these two fleras have competed for the occupa-
tion of these two habitats, and the way in which they have reacted to the
environmental conditions.

Superficial survey would suggest that in a. region such as this with alternating
basalt and sandstone, the geological formation must be the habitat factor of most
outstanding significance; yet observations have disclosed the striking fact that
distribution of the communities is not to be correlated with this but rather with
the edaphic water supply. Apart from the role of humus in the determination of
soil moisture-content, neither chemical nor physico-chemical properties of the soil

BY J. MCLUCKIE AND A. H. K. PETRIE. ' 183

appear to have any fundamental distributional significance, except in the case of
several minor subordinate species. Given the climate and the moderately favour-
able summer rainfall such as they are, wherever the aspect is sheltered, and
conditions, whether they be due to drainage or to the physical structure of the
soil, permit the establishment of a permanent high edaphic moisture-content, Rain-
Forest holds its own; wherever, on the contrary, the true climatic climax habitat
conditions establish themselves on the dry and bleak sandstone plateau, the
sclerophyllous Hucalyptus Forest has unchallenged sway; and wherever conditions
are of an intermediate nature, transitional types of community are to be found.

As progression or retrogression appears to be the fundamental feature in the
development of plant communities we regard the Rain-Forest as a vestigial
remnant of a former vegetation of widespread range in eastern Australia, which,
now faced in most parts with xerophytie conditions inimical to its survival, has
retreated to such habitats as we have shown still to furnish it with optimum
conditions. The xerophytic habitats provided by the elevation of the Blue Mountain
Plateau of sandstone, favoured the advance of the xerophilous endemic Flora which
now occupies the greater part of this formerly Rain-Forest-clad terrain. Yet,
though the old Malayan Flora is securely entrenched in the most favourable
mesophytic habitat at Mount Wilson, and though it can even displace the
autochthonous element in changing habitats, still wherever the conditions become
a little more extreme, Hucalyptus secures a footing; and, as through future ages
the basalt hills are slowly carried away by the unceasing agents of geological
denudation, the area of the Malayan Forest will shrink before the dominance of
the autochthonous Flora.

Summary.

1. This paper is a study of the habitat factors, and an attempt to correlate
these with the interesting distributional features of the vegetation of Mount
Wilson.

2. An explanation of the small amount of invasion of the basalt by types from
the Hucalyptus forests of the sandstone, and why the Rain-Forest grows as well
in many sandstone gullies as on the basalt caps is given in this paper.

3. An analysis of the data of rainfall, temperature, humidity and evaporative
powers of the atmosphere in the different habitats, of the hydrogen-ion concen-
tration of the soil solution, of the moisture-content of the soil is made, and the
relative significance of these factors upon the distribution and organization of
the various plant communities is discussed.

4. The Py value of the soil solution is proved to be an insignificant factor
in the distribution of the vegetation at Mount Wilson.

5. The moisture content of the soil—affected by its humus content, by the
component clay fraction, by drainage, exposure ete, is of the greatest
significance as a controlling factor of the distribution throughout this area. The
significance of aspect and moisture-content of the soil outweigh the effect of all
other factors.

6. The response of the more important components to the habitat factors is
discussed; the general development of the root system in the various habitats,
and the structure of the photosynthetic organs are investigated and correlated
with the environment.

7. The distribution of the communities throughout the Mount Wilson area

is controlled by the edaphic water supply rather than by the nature of the
geological formation.

184 AN ECOLOGICAL STUDY OF THE FLORA OF MOUNT WILSON, iv.

8. The Ceratopetalum-Doryphora association is the dominant community in
the most favoured habitats, e.g. on the basalt slopes, and in the sandstone gullies.

The sclerophyllous Eucalyptus forests, represented by the FHucalyptus
goniocalyz-E. Blaxlandi and the Eucalyptus piperita-Eucalyptus haemastoma asso-
ciations, are the climax communities wherever the environmental factors fall below
the optimum required for the Ceratopetalum-Doryphora association, e.g. on the
sandstone slopes and more exposed habitats of the area.

9. The Malayan exotic flora represented by the Ceratopetalum-Doryphora
association is a mesophytic community which formerly had a very extensive
distribution in New South Wales but is now preserved in sheltered valleys and
on basalt residuals. Its present area will become smaller as the agents of
denudation carry away the basalt caps of the sandstone.

Addendum.

Note on the occurrence of Fucalyptus amygdalina var. nitida.

Since the publication of the previous parts of this work Hucalyptus amygdalina
Labill. var. nitida Benth. has been observed at Mount Wilson. It was found as an
almost pure consociation of the Hucalyptus goniocalyx-H. Blaxlandi association on a
south-westerly slope. H. Blaxlandi was occasional, and the usual stratum-societies
of Alsophila and Pteridium were present.

References.

BroucH, P., McLuckisz£, J., and PrEtTrig, A. H. K., 1924.—An Ecological Study of the
Flora of Mount Wilson. Part i. The Vegetation of the Basalt. Proc. LINN. Soc.
IN-SIW)., xtix, p. 475.

CANNON, W. A., 1915.—The Relation of Root Growth and Development to the Temperature
and Aeration of the Soil Amer. Jour. Bot. 2.

, 1924.—General and Physiological Features of the Vegetation of the more Arid
Portions of Southern Africa, with Notes on the Climatic Environment. Carneg.
Inst. Wash. Publ. 354. \

CLEMENTS, F. E., 1916.—Plant Succession. Carneg. Inst. Wash. Publ. 242.

McLuckKIE, J., and PETRIE, A. H. K., 1926.—An Ecological Study of the Flora of Mount
Wilson. Part iii. The Vegetation of the Valleys. Proc. Linn. Soc. N.S.W., li, p. 94.

PETRIE, A. H. K., 1925.—Id. Part ii. The Eucalyptus Forests. Proc. LINN. Soc. N.S.W.,
horas ales.

RUSSELL, E. J., 1921.—Soil Conditions and Plant Growth.

SHREVE, F., 1914.—A Montane Rain-Forest. Carneg. Inst. Wash. Publ. 199.

WHERRY, E. T., 1920.—Soil Acidity and a Field Method for its Measurement. Ecology, i.

, 1924.—How much Water should be added in making Determinations of the Soil
Reaction?’ Ecology, v, p. 310.

TWO NEW SPECIES OF SETARIA FROM WESTERN AUSTRALIA.

By Dr. A. S. HircHcock, Systematic Agrostologist of the United
States Department of Agriculture.

(Communicated by W. M. Carne.)
[Read 27th April, 1927.]

Recently I received from Mr. W. M. Carne, Botanist and Plant Pathologist
of Western Australia, two species of Setaria which appear to be undescribed. At
the request of Mr. Carne I am submitting technical descriptions. It is unfortunate
that there is not more material at hand, but it will aid in the study of this
critical genus if these tentative descriptions are placed on record.

SETARIA BUCHANANI, Nn. SD.

Planta annua; culmis glabris, ad 70 cm. altis; laminis planis, scabris, 15-25 cm.
longis, 2-6 mm. latis; panicula densa, spiciformis, basi interrupta, 7-12 cm. longa;
axi scabro; ramulis 1-1-5 cm. longis, suberectis; setis antrorsum scabris; spiculis

ellipticis, 24 mm. longis; gluma prima spiculam 4 aequante, acuta; gluma secunda
' longitudine spiculae; lemmate sterile longitudine spiculae: lemmate fertile
rugulose. |

Annual; culms branching at base, glabrous (scabrous below panicle), about
70 em. tall; sheaths glabrous; ligule 1 to 1-5 mm. long, membranaceous at base,
Dilose above; blades flat, scabrous, 15 to 25 cm. long, 2 to 6 mm. wide; panicle
dense, spikelike, pale green, interrupted, 7 to 12 cm. long, the axis scabrous,
angled, the branches 1 to 1:5 cm. long; bristles solitary below a part of the
spikelet, antrorsely scabrous, 2 to 3 times as long as the spikelets; spikelets
elliptic, glabrous, about 2-4 mm. long; first glume about half as long as the
spikelet, acute; second glume as long as spikelet, 5-nerved; sterile lemma as long
as the spikelet, 5-nerved, with an additional pair of nerves above; fertile lemma
about as long as the sterile lemma, rather faintly transversely rugulose, pale.

Type specimen: Murchison District, Western Australia, G. Buchanan, H. 201,
April, 1925.

This species differs from Setaria viridis (L.) Beauv. in the elongate blades,
the interrupted panicle, the fewer bristles, and the longer acute first glume.

SETARIA CARNEI, DN. Sp.

Planta perennis; culmis glabris, 40-50 cm. altis; laminis planis, scabris,, 10-20
em. longis, 2-5 mm. latis; panicula densa, spiciformis, interrupta, 7-12 cm. longa;
axi scabro; setis retrorsum scabris; spiculis ellipticis, glabris, 2 mm. latis.

Perennial; culms glabrous (roughish below panicle), 40 to 50 cm. tall;
sheaths smooth, compressed; ligule about 1 mm. long, pilose; blades flat, glabrous
beneath, scabrous on the upper surface, 10 to 20 cm. long, 2 to 5 mm. wide; panicle

dense, spikelike, interrupted, pale, 7 to 12 cm. long, the axis scabrous, angled, the
X

186 NEW SETARIA FROM WESTERN AUSTRALIA.

branches 3 to 5 mm. long; bristles solitary below each spikelet, retrorsely scabrous,
2 to 4 times as long as the spikelet; spikelets elliptic, glabrous, 2 mm. long; first
glume about half as long as the spikelet, acutish; second glume as long as the
spikelet, distinctly 7-nerved; sterile lemma as long as the spikelet, 5-nerved, with
an additional pair of nerves above; fertile lemma rather faintly transversely
rugulose, pale.

Type specimen: Broome, Western Australia, collected by the North West
Department, H. 197, April, 1925. In the U.S. National Herbarium there is a
specimen of this species from West Australia, collected by Drummond.

This species resembles S. buchanani in aspect but differs in being apparently
perennial, in the retrorsely scabrous bristles, and in the smaller spikelets.

aoe
a A heer ce OR
aia eke Fad) ek
Pie BURL AY

vey

Proc. Linn. Soc. N.S.W., 1927. PLATE I.

; Li hy.
4 “cl i

=“ To DUNGOG

N = 205° |
4} v t\ONBLAaTEAUASe ON l atl" ak f
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10400

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To Y
SEAHAM

GEOLOGICAL MAP

OF” ThE
LAMB’S VALLEY-PATERSON
DISTRICT

* To STANHOPE

SCALE

To DALWOOD

SALES,

id

i

Wigee sot

_ a | ys

, op oF A AoA
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Proc. Linn. Soc. N.S.W., 1927. PLATE Il.

1,2. Deltopecten rienitsi, n. sp. 3-5. Estheria coghlani Cox.
6. E. glenleensis, n. sp. 7, 8. H. wianamattensis, n. sp.

Proc. Linn. Soc. N.S.W., 1927. PLATE Itt.

1-4. Estheria ipsviciensis, n. sp. 5, 6. EH. novocastrensis, N. Sp.
7. E. lenticularis, n. sp. 8, 9. H. lata, n. sp.

Proc. Linn. Sou. N.S.W., 1927. PLATE IV.

1. EHstheria obliqua, n. sp. 2, 3. E. glabra, n. sp. 4. H. linguiformis, n. sp.
5. E. belmontensis, n. sp. 6. EH. trigonellaris, n. sp.
7, 8. E. (?) bellambiensis, n. sp.

Proc. Linn. Soc. N.S.W., 1927. PLATE V.

oe ee ci

Photomicrographs of fossil wood from Ulladulla, N.S.W.

Proc. Linn. Soc. N.S.W., 1927. M PLATE VI.

Photomicrographs of fossil wood from Ulladulla, N.S.W.

Proc. Linn. Soc. N.S.W., 1927. PLATE VII.

1. Grevillea laurifolia. 2-5. Forms of G. Gaudichaudii.
6. G. acanthifolia.

‘

ay

a
Ny
4

Proc. Linn. Soc. N.S.W., 1927. PLATE VIII.

1. Character of deposits of fiood silts.

2. Cracks due to drying in flood silts.

a.

' ‘
m 7 Ley A
ics 0 Newer iw lyes
. ou rans 2
Le ;
;

aa
> *

Proc. Linn. Sou. N.S.W., 1927. PLATE [X.

1347. Galls of Kladothrips augonusaxxos, n. sp. on Acacia doratoxylon.
518. Galls of Choleothrips geijerae, n. sp. and Dolerothrhips geijerae, n. sp.
on Geijera parviflora.

THE VEGETATION OF THE KOSCIUSKO PLATEAU.
Part i. THE PLANT COMMUNITIES.

By JoHn McLucxigr, M.A., D.Sc., Assistant-Professor of Botany and
Lecturer in Plant Physiology and Ecology,
the University of Sydney;
AND -
ArtHuUR H. K. Petrig, M.Se., Senior Demonstrator in Botany,
the University of Melbourne.
(From the Botanical Laboratory, University of Sydney.)

(Plates x-xix and eight Text-figures. )
[Read 27th July, 1927.]

Contents.

Introduction.
Meteorological Data.
The Hucalyptus Forests.

The Eucalyptus coriacea Consociation.

The Eucalyptus Gunnii Consociation.

The Hucalyptus coriacea-E. Gunnii Ecotone and the EH. stellulata Consociation.

The Leptospermum Shrub Society.

Regeneration of the Hucalyptus Forests after Fire.

Factors causing or modifying the Timber-line.
The Marsh Vegetation.

The Poa-Hypolaena Ecotone.

The Hypolaena Associes.

The Restio Associes.

The Richea Associes.

The Epacris Associes.

The Baeckea-Callistemon Association.

The Luzula-Carex Associes.
The Alpine Vegetation.

The Poa-Celmisia Association.

The Epacris petrophila Consociation
Summary.

Introduction.

The Kosciusko Plateau, the plant ecology of which forms the subject of this
paper, lies in the Snowy Range in the southern portion of New South Wales,
and is the highest land surface on the Continent.’ It is an elevated block about
20 miles long by 6 miles broad, rising gradually from east to west. At the
junction of the Thredbo and Snowy Rivers, it is 3,000 feet above the sea. Mount
Kosciusko at the other extreme (its westerly limit) is about 7,328 feet high.
Only a few isolated points rise above 7,000 feet. A horse shoe-shaped area
surrounding the Snowy head-waters exceeds 6,500 feet (contour map Plate x? and
block diagram Text-fig. 1).

1The following description of the topography of the Kosciusko Plateau is taken from
the paper “The Kosciusko Plateau,” by G. Taylor, W. R. Browne and F. Jardine,
published in the Proc. Roy. Soc. N.S.W., 1925, pp. 200-205.
2The block, from which this map has been printed, has been kindly lent by the
Royal Society of New South Wales, it having appeared in the Proceedings of that Society
for 1925 as Plate i.
A

N OF THE KOSCIUSKO PLATEAU, i,

EGETATIO

THE V

188

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BY J. MCLUCKIE AND A. H.’K. PETRIE. 189

“There is a very marked drop in the Plateau east of Perisher Creek, for
practically all the high land (over 6,000 feet) lies to the west”. . . “The
5,000 feet contour bounds the Plateau uplands around the lower Snowy River,
which flows through a gorge which is largely below that level’. “The drainage
is arranged according to a very interesting plan. The main stream of the
district is really the broad Eucumbene Valley in the extreme east. This is
longer and ‘older’ than the Snowy Valley. The latter has cut back as a gorge into
the uplifted plateau—probably along fault planes—as far back as Charlotte’s Pass.
Above this the valley is fairly wide and approaches maturity. Further west, the
head-waters, above the road-ford, flow and meander through a flat senile valley.
The Crackenback-Thredbo Valley is a profound gorge for most of its extent.

5 Besides these two parallel gorges of the Snowy and Crackenback, there
is a third more or less continuous valley running between: them. . . . It
consists of a series of shallow basins linked by cols or gaps. . . . Hach of
these basins drains to the north by a creek leading into the Snowy River: This
‘Road Valley’ is about 1,000 feet above the two parallel river valleys. Its
topography is probably in part due to bygone giacial erosion”.

This region is the coldest portion of Australia, and for six months in the
year the higher elevations are covered in snow. It thus offers features of special
interest so far as the vegetation of Australia is concerned. The Plateau is an
elevated land surface composed of gneissic granite, which ascends abruptly from
the surrounding plains at about 3,000 feet above sea-level, to an altitude of 5,000
to 6,000 feet, from which level it rises more gradually until it culminates in
various peaks ranging up to 7,328 feet. The lower slopes are steep and well
drained, and are dissected by various defiles formed by erosion; the higher
portion of the Plateau, however, above 5,000 feet, is traversed by long shallow
valleys, whose flat bottoms form marshes by the slow draining away of the water
from the melting snow of the higher peaks.

A consideration of the plant communities has led us to recognize three unit-
areas in the region under discussion:

(1) The montane zone, from 3,000 to approximately 5,000 feet, comprising
the lower slopes of the Plateau.

(2) The sub-alpine zone, from approximately 5,000 feet to the tree-line at
6,000 to 6,500 feet.

(3) The alpine zone, from the tree-line to the highest elevations.

The montane zone is entirely clad with sclerophyllous Eucalyptus forest,
as is characteristically the case with the mountain regions of Eastern Australia.
Eucalyptus forest also clothes the ridges and slopes in the sub-alpine zone; the
boundary between the montane and sub-alpine zones, however, is also the position
of the junction of the two main consociations of the forest. The lower portions
of the slopes of the sub-alpine region, below the forest, are occupied by low-
tussock grassland, which, admixed with mat herbage, is also the main formation
of the alpine zone. A marsh vegetation occupies the shallow valley-bottoms
throughout the sub-alpine and alpine regions. These three types of vegetation
will be discussed in detail in the subsequent portion of the paper.

The lists of plants from this area, published by Maiden (1898-99) and Helms
(1896-97) proved of great value to the writers in their identification work, but did

not furnish an analysis of the communities of the plateau, which is attempted in
this paper.

190 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

Meteorological Data.

The following information dealing with the distribution of snow falls
throughout the season, the mean monthly humidity and the absolute maximum
and minimum temperatures, has been kindly supplied to us by Mr. D. J. Mares,
Divisional Meteorologist of the Commonwealth of Australia.

The greatest mean monthly humidity occurs during tne winter months from
about May to September, during which period considerable snow or rain falls,
and heavy frosts are frequent.

The absolute maximum temperature so far recorded is 88° on the 9th November,
1919. The. mean monthly humidity is lowest during the summer months.

The lowest absolute minimum temperature for any month occurred on the
15th August, 1920, and other very low temperatures during 1919 in the months
July to September of the same and the following years, that is just subsequent
to the devastating fires which. swept over the plateau and destroyed great areas
of the Poa-Celmisia and Eucalyptus coriacea-E. Gunnii associations, and the
#H. stellulata consociation. Perhaps the severity of the climate during such
periods has been responsible for the tardy renaissance of the Zucalyptus forests
and for the killing of areas near the highest portion of the tree-line.

Low temperatures and severe frosts during certain winters if followed by
droughty conditions during the summers, retard the development of buds, and
flowering and fruiting, and may temporarily destroy the more exposed parts of
the Poa and Eucalyptus associations.

' Commonwealth Meteorology.
Mt. Kosctusko.—-TABLE I.

Mean Monthly Humidity 9 a.m. (14 years).

Jan. Feb. | Mar.| April} May | June| July |. Aug.| Sept.| Oct. Nov. | Dec.

56% | 60% | 62% | 65% 69% | 79% | 84% | 79% | 70% | 60% | 55% | 58%

Absolute Maximum Temperature, 1911-24 (Fahrenheit).

Jan. | Feb. | Mar.| April| May | June | July | Aug.} Sept. | Oct. Nov. | Dec.

84-0 85-0 77-0 71-0 67-0 56-0 52-2 57-1 78-0 78-0 88-0 | 78-0
30th | 15th 5th 4th 2nd 7th 25th | 27th |} 26th} 10th 9th 9th

10th

1912 | 1919 | 1924 | 1911 | 1922 | 1919 | 1916 | 1914 | 1922 | 1922 | 1919 | 1918
‘ 4th 29th 1st
1923 1924 1924

Absolute Minimum Temperature, 1911-24.

| | |
Jan. | Feb. Mar. April | May June | July een Sept. Oct. Nov. | Dec.
i lentes ace aes A
21-0 | 24-9 | 19-8 | 15-0 |) 12:00) Wao 70 6-3 | 14:0 | 16:0 | 13-0 | 20-0
27th 5th) 24th | 27th | 28th | 22nd | 16th 15th 8th 22nd 4th 3rd
POLS LST LOL, Hat | 1913 | 1917 | 1919 | 1920 | 1919 1918 | 1911 | 1924
191 | |

BY J. MCLUCKIE AND A. H. K. PETRIE: 191

Mt. Kosciusko.—TABLE II.
Days of Snow.

Jan. Feb. | Mar.} April| May | June | July | Aug.| Sept. |} Oct Nov. | Dec.
1924 | 1924 | 1924 | 1924 | 1924 | 1924 | 1924 | 1924 | 1924 | 1924 | 1924 | 1924
—t | 1 — 2 2 37 4 3t 7 — — —

* Snow still abundant round summit of Mt. Kosciusko.

+ Frosts more severe and sustained than for several years.

t Rarity remarkable. Frosts severe.

Jan. | Feb. | Mar.| April| May | June July | Aug.| Sept. | Oct. Nov.| Dec.
1923 | 1923 | 1928 | 1923 | 1923 | 1923 | 1923 | 1923 | 1923 | 1923 | 1923 | 1923
iL — — — | 4 5 §* 2 5 1 — —

* So much snow over such an extended area not seen since 1914.

Jan. Feb. | Mar.| April| May | June July | Aug.| Sept. | Oct. Nov Dec.
1922 | 1922 | 1922 | 1922 | 1922 | 1922 | 1922 | 1922 | 1922 | 1922 | 1922 | 1922
—_— oe 1 1 2 11 6 aba ook 6 4 — —
Jan. Feb. | Mar.| April| May | June | July | Aug.| Sept. Oct. Nov. | Dec.
1921 | 7921 | W921 | L92T | L921 | 1921) L921 7 1921 | W921 | 1921 | 1927 |) 1921
ae poe — = 2 ons 5T 9 1 2 — —

* Season mild. Very little snow below 6,000 feet level.

+ More rain than snow.

THE EUCALYPTUS FORESTS.

In the region under study there is only one association of Hucalyptus forest,
namely, the Hucalyptus coriacea-E. Gunnii association, which extends from the
foot of the Snowy Mountains on the edge of the Monaro Plains up to the tree-line.
It occurs also on the exposed wind-swept Monaro Plains themselves, from”

which the Snowy Mountains rise.
The following three consociations occur:

1. The Eucalyptus coriacea consociation, which is typical of the sub-alpine
zone, and is found chiefly from about 4,500 feet to the tree-line between 6,100 and

6,500 feet, although it occurs also at lower altitudes.

2. The Eucalyptus Gunnii consociation which is found chiefly between 3,500
and 5,000 feet, and generally prefers the more sheltered slopes characteristic of

the montane region.

3. The Eucalyptus stellulata consociation which occupies small areas between

the altitudes of 4,000 and 5,000 feet.

THE EUCALYPTUS CORIACEA CONSOCIATION.

Habitat.

While the main adaptive feature of Hucalyptus forest is xerophily,

the

Eucalyptus coriacea consociation has fitted itself also for existence amid the

192 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

severities of a sub-alpine climate, and is the only forest which has ascended above >
an altitude of 5,000 feet in Australia. The same community occurs on the Monaro
Plains, at an altitude of 2,700 to 4,000 feet, where it-is exposed to bleak and
desiccating winds from the west. Hucalyptus coriacea has also been recorded
from other mountain tops in New South Wales, where it is found in similar
habitats: aura

So. far as the Snowy Mountains are concerned, the Eucalyptus coriacea
‘consociation is found chiefly from about 4,500 to 5,000 feet up to 6,500 feet, which
is the absolute limit of tree vegetation. It probably occurs, however, at a much
lower altitude than 4,500 feet on the western slopes of the mountains owing to
exposure to west and north-west winds. Above 5,000 feet the forest is confined
to the upper and middle portions of the slopes of the hills, giving place below to
grassland (Plate xi, figs. 1 and 2; Plate xii, fig. 3). All flat and gently sloping
ground above this altitude is extremely damp, or even water-logged, owing to the
drainage from the melting snow on the upper slopes, combined with the impervious
nature of the granite substratum. The restricted distribution of the Hucalyptus
forest which avoids all swampy areas is probably the result of the water-logging
and consequent lack of aeration of the lower strata of the soil. Near the tree-
line the forest is still further limited on account of the greater amount of snow,
and the trees are confined to the drier ridges (Plate xvi, fig. 4).

Structure and Physiognomy.

Eucalyptus coriacea is not only dominant, but is the only tree in the
consociation at the higher altitudes; it has a distinctive appearance imparted by its
smooth white bark, gnarled branches, excessively glaucous twigs, and thick
coriaceous leaves. At an altitude of about 5,000 feet, in less exposed areas,
Hucalyptus coriacea attains a height of about fifty feet and forms an extensive
and fairly close forest (Plate xi, fig. 1); at higher altitudes, however, and on the
hill-tops above 5,000 feet, the trees become stunted and the forest is more open
(Plate xii, fig. 3); while near the tree-line they are rarely taller than twelve to
fifteen feet, and often their gnarled branches and trunks are almost prostrate
(Plate xvi, fig. 4). This condition is no doubt the result of the heavy snow-falls
and frosts during several months of the year and the stimulation of excessive
transpiration by strong winds, which are frequent at these altitudes.

While Eucalyptus coriacea forms a pure consociation at the higher altitudes,
£. stellulata becomes frequent at 5,000 feet, as will be described subsequently.
E. Gunnii is occasional at 5,000 feet, and also becomes abundant in Digger’s Creek
valley, where an extensive ecotone region exists between the £. coriacea and
E. Gunnii consociations. EH. viminalis is also occasional in this ecotone region;
since, however, the lower strata are highly typical of the H. Gunnii consociation,
neither this species nor EH. viminalis can be considered typical of the HH. coriacea
forest.

Shrubs form a conspicuous element in the Hucalyptus forests of certain slopes,
occurring in dense societies composed chiefly of a dominant species, or else
scattered and mixed in composition. In more exposed areas, e.g., on the top of
the ridges, shrubs are absent, and the ground is occupied by Poa caespitosa,
forming low tussocks, and by other grasses and herbaceous types, which also
occupy the ground between the shrubs when they are so scattered as to permit
of the development of lower strata.

Between 5,000 and 5,500 feet societies of Oxylobium ellipticum var. alpinum are
common, containing as subordinates Pimelea ligustrina, Drimys aromatica var.
pedunculata, Cassinia aculeata, Olearia myrsinoides, Pimelea axiflora and Veronica
Derwentia,

40 Feet

UPPER

# SS 5
*ue sss
s

Ss
Sis

BY J. MCLUCKIE AND A. H. K.

17 Feet

PETRIE. 193

Text-figure 2, Chart of an
area in an. open Hucalyptus
coriacea forest (5,500 ft.).

This chart illustrates the
vegetation on a typical slope.
The lower end of this chart
is within a few feet of a small
creek traversing the hill-side.

As_ the water is approached
certain types such as Oreomyr-

rhis andicola and Veronica
disappear, while species. of
Helichrysum become more
abundant.

Veronica is always abundant
on the slopes of the water
courses, while Oxylobium
ellipticum prefers the drier hill-
sides more remote from the
streams.

@ ~ Veronica Derwentia
(0%- Oxylobium ellipticum.

S @=Stellarium pungens.

H = Helichrysum bracteatum.
G@ — Geranium dissectum..

T = Tarazacum sp.

Or- Oreomyrrhis andicola.
Hs = Helichrysum scorpioides.
Ep ~ Epilobium glabellum.

Se - Senecio pectinatus.

Ol - Olearia myrsinoides.

&9)- Eucalyptus coriacea, seedlings.

Ca - Cassinia aculeata.

All space between symbols is
occupied by Poa _ caespitosa
forming a close grassy cover-
ing with occasional tussocks and
withered patches. An attempt
has been made to convey an
impression of the relative pro-
portion of the area that is
occupied by each component.

Scale: 1 cm. = 2 feet.

194 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

The Oxylobium society is replaced by one dominated by Callistemon Sieberi in
the little watercourses which frequently run down the slopes to the creeks and
swampy areas invariably occupying the flat bottoms of the shallow valleys. This
society is closely related to the Callistemon Sieberi consociation of the marsh

vegetation, and contains a number of swamp-loving types not found elsewhere in
the Eucalyptus forests.

Floristic Composition.
Shrub Stratum

Callistemon Sieberi D.C. d*
Cassinia aculeata R.Br. f
Oxzylobium ellipticum R.Br., var. alpinum

Maiden and Betche o-f
Drimys aromatica F.v.M., var. pedunculata

Maiden co)
Panax sambucifolius Sieb. r
Leptospermum lanigerum Sm. If

Ground Stratum
Richea Gunnii For.
Acaena sanguisorba Vahl.
Anagallis arvensis Linn.
Epilobium glabellum Forst.
Rumeax acetosella Linn.
Taraxacum dens-leonis Desf.
Stylidium graminifolium Swartz.
Linum marginale Cunn.
Blechnum penna-marina (Poir.) Kuhn
Galium umbrosum Sol.
Sphagnum societies
Prostanthera cuneata Benth.
Epacris paludosa R.Br.

ii ©) ©) ©C) 6) © ©) ©. ©) fs eo !n0

=
e}

Veronica Derwentia also forms dense societies (Text-fig. 2), especially in more
open areas, with Olearia megalophylla (f{), Drimys (f), and Ozylobium ellipticum
var. alpinum (o) (Plate xiii, fig. 2).

Above 5,500 feet these societies are not so frequent, their place being taken
by Prostanthera cuneata, which forms abundant societies up to and beyond the
tree-line (Plate xvi, fig. 4; Plate xix, fig. 2). Phebalium ovatifolium is a frequent
subordinate in this society. Elsewhere the shrubs are mixed or absent, Helichrysum
baccharoides and H. rosmarinifolium being conspicuous.

On the tops of the ridges, and between the shrubs where they are less dense,
Poa caespitosa, with its many herbaceous subordinates, forms a close covering.
At the foot of the slopes and at the tree-line this stratum-society passes out
unchanged beyond the trees, and assumes the rank of an association, a feature
which will be referred to again subsequently (Plate xii, fig. 3).

Floristic Composition.
Tree Stratum.

Bucalyptus coriacea Cunn. d
E. stellulata Sieb. f (up to 5,000 ft.)—o (to 5,500 ft.)
E. Gunnii Hook. o (to 5,000 ft.)
Tall-Shrub Stratum.
Helichrysum rosmarinifolium Less. f-c (above 5,500 ft.)
Drimys aromatica F.v.M. var. pedunculata Maiden f-la
Cassinia aculeata R.Br. f-la
Veronica Derwentia Littlej. f-la

*In these lists, a = abundant, ec = common, d = dominant, f = frequent, 1 = locally,
o = occasional, r = rare, sd = subdominant, vr = very rare.

Tall-Shrub Stratum.

Hovea longifolia R.Br.

Oxylobium ellipticum

BY J. MCLUCKIE AND A. H. K. PETRIE.

R.Br. var. alpinum Maiden

o-a (5,200-6,000 ft.)

and Betche
Bossiaea foliosa Cunn. o-la
Olearia myrsinoides F.v.M. o-le
Helichrysum baccharoides F.v.M. o-le
Olearia megalophylla F.v.M. o-lf-le
Pimetlea ligustrina Labill. var. hypericina Benth. o-f

Phebalium ovatifolium F.v.M.
Pimelea axviflora F.v.M.

Rubus rosifolius Sm.

Orites lancifolia F.v.M.
Grevillea australis R.
Hakea Macreana F.v.
Lissanthe montana R.Br.

Acacia alpina F.v.M.
Panax sambucifolius

Prostanthera cuneata Benth.

fo)
ro)
{o)
Br. Oo
M. Ts
r
Tt
Sieb. vr (below 5,100 ft.)

Leptospermum lanigerum Sm. if

Callistemon Sieberi D.C.

Low-Shrub Stratum.

Hibbertia linearis R.

Ground Stratum.

Poa caespitosa Forst. a-lo
Taraxacum dens-leonis Desf. f-a
Trifolium sp. f-a
Rumex acetosella Linn. ; f-la
Helichrysum scorpioides Labill. f-¢
Geranium dissectum Linn. fy
Ranunculus lappaceus Sm. ifs
Plantago varia R.Br. f
Senecio lautus Sol. f
Wahlenbergia gracilis D.C. if
Acaena sanguisorbae Vahl. o-a
Stellaria pungens Brongn. o-a
Brachycome scapiformis D.C. , o-a
Craspedia Richea Cass. o-a
Podolepis longipedata Cunn. o-la
Stylidium graminifolium Swartz. o-f
Epilobium glabellum Forst. {o)
Viola betonicifolia Sm. fo)
Scaevola Hookeri F.v.M. societies. °
Comesperma retusum Labill. Oo (5,000 ft.)
Brachycome decipiens Hook. Oo
Poranthera microphylla Brongn. ra)
FPolystichum aculeatum (l.) Schott. o (under rocks)
Kunzea Muelleri Benth. fo}
Celmisia longifolia Cass. o-f
Stellaria flaccida Hook. {o)
Oreomyrrhis andicola Endl. ro)
Euphrasia Brownii F.v.M. fo)
Helichrysum bracteatum Willd. (0)
Blechnum penna-marina (Poir.) Kuhn. r
Galium umbrosum Sol. r
Linum marginale Cunn. ie
Dianella tasmanica Hook. r
Podocarpus alpina R.Br. societies r (above 5,600 ft.)

Asplenium flabellifolium Cav.

Chiloglottis Gunnii Lindl. Ic
Pterostylis mutica R.Br. lf
Richea Gunnii For. lo
Sphagnum societies lo
Anagallis arvensis Linn. lo (5,000 ft.)

Br. var. obtusifolia Benth. 0 (below 5,000 ft.)

Epacris paludosa R.Br. lo

f (5,500 ft. upwards)

o-f (above 5,600 ft.)

(near tree-line)
(at tree-line)

vr (5,300 ft.)—a (6,000 ft.)

lf (5,000 ft.)

(above 5,200 ft.)

vr (5,000 ft. downwards)

195

196 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

THE EUCALYPTUS GUNNII CONSOCIATION.

Passing into the sheltered gullies just below 5,000 feet, the Eucalyptus coriacea
consociation gives place to the H. Gunnii consociation, very gradually in Digger’s
Creek valley, comparatively abruptly in other areas. H. Gunnii grows taller than
E. coriacea, the trees averaging about seventy feet in height. For some distance
into this consociation a number of shrubs characteristic of the H. coriacea forest
persist, but a distinct and continuous change is noticeable as one descends the
Plateau, new types appearing and former ones disappearing. Poa caespitosa
tussocks are much less abundant; although they still form a carpet in certain
areas where the soil is damper, a considerable amount of the ground is quite bare
of grass or only sparsely covered (Plate xvi, fig. 3). In general this forest has a
more xerophilous impress since many of the herbaceous types of the H. coriacea
consociation are absent, the rich carpet of grass is less evident, and the shrubs
have a more scraggy, sclerophyllous appearance; indeed the general physiognomy
of the community resembles that of the H. piperita forest characteristic of the
sandstone plateau at Mount Wilson.

The smaller rainfall which characterizes the lower altitudes of the mountains
probably contributes to this feature of the ZH. Gunnii consociation, as also does a
change in the topography. We are here below the level of the marshy flats and
the continual streams of water from the melting snow; the water is concentrated
in the deep gorges of the Snowy River and its tributaries, by means of which the
gravitational water passes rapidly away, resulting in a general lowering of the
soil moisture-content. Moreover this consociation dominates an area from which
the snow disappears early on the approach of summer; there is therefore less soil
moisture as compared with the higher zones, and the plant life is dependent directly
upon the rainfall for its moisture requirements throughout the summer months.
The extremely xerophilous structure of the plants inhabiting this zone indicates an
approximation t6 the conditions which have stimulated the development of the
highly xerophytic sclerophyllous forest on the Hawkesbury Sandstone habitat
around Sydney.

Floristic Composition.~

Tree Stratum.

Eucalyptus Guninii Hook. d

E£. viminalis Labill. o-c

E. coriacea Cunn. o-f

E. stellulata Sieb. fo)

E. amygdalina Labill. var. nitida Benth. Ts
Low-Tree Stratum.

Acacia rubida Cunn.
Shrub Stratum.

Ozylobium ellipticum R.Br. var. alpinum Maiden

and Betche

Bossiaea foliosa Cunn.

Daviesia ulicina Sm.

Leptospermum stellatum Cay.

Cassinia aculeata R.Br.

Baeckea Gunniana Schau.

Grevillea lanigera Cunn.

Lomatia longifolia R.Br.

Acacia penninervis Sieb.

Rubus parvifolivs Linn.

Panag sambucifolius Sieb.

Daviesia corymbosa Sm.

o}

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ei
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= OL OTOUO FOL OROTO ES
mh ho

BY J. MCLUCKIE AND A. H. K. PETRIE: 197

Low-Shrub Stratum.
Discaria australis Hook. (0)

Grass Stratum.
Poa caespitosa Forst. c-O
Wahlenbergia gracilis D.C. f-c
Stellaria pungens Brongn. f-c
Taraxacum dens-leonis Desf. f-c
Hibbertia linearis R.Br. var. obtusifolia Benth. f-c
Helichrysum semipapposum D.C.
Geranium dissectum Linn.
Rumex acetosella Linn.
Brachycome scapiformis D.C.
Plantago varia R.Br.
Epilobium glabellum Forst.
Trifolium sp.
Velleya paradoxa R.Br.
Acaena sanguisorbae Vahl.
Prunella vulgaris D.C.
Xerotes longifolia R.Br.
Pimelea glauca R.Br.
Arthropodium paniculatum R.Br.
Persoonia chamaepeuce Lhotsky
Stylidium graminifolium Swartz.
Galium umbrosum Sol.
Lotus australis Andr.
Lotus corniculatus Linn.
Helichrysum scorpioides Labill.
Pterostylis coccinea Fitzg. vr
Bulbine bulbosa Haw. vr
Erythraea australis R.Br. pala}

fini
LS (a)
i)

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fear) Lar) ta
©

OOO © OO © OOO [ay trp Inoilnd [nd tnd 1d

THE EUCALYPTUS CORIACEA—E. GUNNII ECOTONE AND THE E. STELLULATA CONSOCIATION.

As has been mentioned earlier, the slopes of Digger’s Creek valley (4,000-5,000
feet) are clothed by a forest which represents an ecotone between the Z. coriacea
and EH. Gunnii consociations. The continuity of this ecotone is interrupted by the
occasional presence of a pure consociation of H. stellulata. Where Digger’s Creek
valley joins the main gorge of the Snowy River (at about 4,000 feet) the ecotone
gives place gradually to the #. Gunnii consociation. One may suppose that in this
region the conditions are intermediate between the exposed habitat yet damp soil
occupied by the E. coriacea forest on the one hand, and the drier, yet more
sheltered environment normally occupied by H. Gunnii. The valley is sheltered,
and the sides are steep; yet the soil in many parts has still much of the dampness
characteristic of the higher altitudes.

We have not up to the present been able to discover definitely the factors
leading to the interruption of the ecotone by occasional consociations of
#H. stellulata, but it is possible that a slightly greater degree of exposure may be
the key to its presence; nor did we observe any difference in the floristic
composition of the two communities.

Floristic Composition.

Tree Stratum. Ecotone. EF. stellulata.
Eucalyptus coriacea Cunn. d fo)
H. Gunnii Hook. a-f —_
EH. viminalis Labill. fo) —_—
EH. stellulata Sieb. to) d

Small-Tree Stratum.
Acacia neriifolia Cunn. o in societies

198 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

Shrub Stratum.
Veronica Derwentia Littlej. f-a
Epacris sp. f
Pimelea axiflora F.v.M. f
Oxylobium ellipticum R.Br. var. alpinum Maiden

and Betche
Bossiaea foliosa Cunn.
Hymenanthera dentata R.Br.
Olearia megalophylla F.v.M.
Hovea longifolia R.Br.
Olearia myrsinoides F.v.M.
Drimys aromatica F.v.M. var. pedunculata Maiden
Pimelea ligustrina Labill. var. hypericina Benth:
Panax sambucifolius Sieb.
Daviesia ulicina Sm.
Acacia decurrens Willd.
Lomatia longifolia R.Br.
Acacia penninervis Sieb.
Persoonia chamaepeuce Lhotsky
Hibbertia linearis R.Br. var. obtusifolia Benth.
Acacia siculiformis Cunn.
Leptospermum stellatum Cav.
Leptospermum lanigerwm Sm.
Callistemon Sieberi D.C.
Hakea Macreana F.v.M.
Rubus parvifoliws Linn.

Low-Shrub Stratum.
Helichrysum semipapposum D.C.
Cassinia aculeata R.Br.

Ground Stratum.
Stellaria pungens Brongn.
Poa caespitosa Forst.
Acaena sanguisorbae Vahl.
Geranium dissectum Linn.
Trifolium sp.
Wahlenbergia gracilis D.C.
Taraxacum dens-leonis Desf.
Galium umbrosum Sol.
Rumex acetosella Linn.
Brachycome scapiformis D.C.
Blantago varia R.Br.
Viola betonicifolia Sm.
Velleya paradoza R.Br.
Velleya montana Hook.
Helichrysum bracteatum Willd
Prunella vulgaris D.C.
Podolepis longipedata Cunn.
Oreomyrrhis andicola Endl.
Linum marginale Cunn.
Craspedia Richea Cass.
Stylidium graminifolium Swartz.
Hypericum japonicum Thunb.
Lotus australis Andr.
Goodenia hederacea Sm.
Poranthera microphylla Brongn.
Xerotes longifolia R.Br.
Helichrysum scorpioides Labill.
Asplenium flabellifolium Cay.
Exzocarpus nana Hook.
Arthropodium paniculatum R.Br,
Pterostylis acuminata R.Br.
Dianella sp.
Gastrodia sesamoides R.Br.

1
Hh

@uOMOMOMmO) C/O} OF OMONOMOMOC Om Gi

(in drainage
channels)

oy

tooo 0

oo

moan0oa0nD

'
ie)

-O
(below 4,650 ft.)

CO) fry tno tnd Ina ind tna) tnd Id

(below 4,400 ft.)
(below 4,100 ft.)

(under rocks)

gpod SE UOT) @)@) © G0) GY QO) -C ©)

BY J. MCLUCKIE AND A. H. K. PETRIE. 199

Ground Stratum. ‘

Stackhousia monogyna Lindl. ? If

Lotus corniculatus Linn. lf-la a Reson
Epilobium glabellum Forst. 1f-le SiEVeae)
Juncus pallidus R.Br. lo J

Lotus australis Andr. ra)

THE LEPTOSPERMUM SHRUB SOCIETY.

Along the banks of the rivers occupying the beds of the valleys whose slopes
are clad by the Eucalyptus Gunnii consociation or by the ZH. coriacea-E. Gunnii
ecotone, occurs a society of shrubs about five to seven feet in height. In the
upper part of the ecotone at about 4,600 feet Leptospermum lanigerum is the
dominant, while, as the beginning of the HE. Gunnii consociation is approached,
Leptospermum stellatum becomes co-dominant. (Plate xiv, fig. 2).

The majority of the components of this society appear to be species requiring
a moderate degree of shelter and soil-moisture. It is interesting to note also that
Kunzea Muelleri and Hakea microcarpa here attain a height of four to five feet,
whereas above the 5,000 feet altitude Kunzea is only about six inches high, and
Hakea one foot. In many cases this society is so dense as to be almost impenetrable
and, during periods of flooding of the river, is practically submerged.

Floristic Composition.
Shrub Stratum, 4,650 ft.
Leptospermum lanigerum Sm.
Callistemon Sieberi D.C.
Daviesia corymbosa Sm.
Leptospermum stellatum Cav.
Cassinia aculeata R.Br.
Hakea Macreana F.v.M.
Kunzea Muelleri Benth.
Hakea microcarpa R.Br.
Correa Lawrenciana Hook.
Lomatia longifolia R.Br.
Ground Stratum.
Juncus pusillus Buch. c
Hypolaena lateriflora Benth. f
Epilobium glabellum Forst. 9)
Lotus australis Andr. (e)
Lotus corniculatus Linn. 6)
Drosera peltata Sm. .
Oreomyrrhis andicola Endl.
Stylidium graminifolium Swartz.
Hypericum Japonicum Thunb.
Utricularia dichotoma Labill. var. wniflora Benth.
Juncus pallidus R.Br.
Creeper.
Glycine clandestina Wendl.

emg
Q a

AO oO Oo: 0 Oo

oo000

ae
rt

fo}

REGENERATION OF THE EUCALYPTUS FORESTS AFTER FIRE.

On account of the high rainfall and general dampness of the ground, bush
fires are by no means so prevalent on the Kosciusko Plateau above an altitude of
5,000 feet as in other more arid regions of the State; and at the time of study
(1924 and 1925) none had occurred since 1919 in the area under review. The fire
which took place in the latter year, however, swept over large areas of the
Eucalyptus coriacea consociation between 5,000 and 5,500 feet, as well as through
a small consociation of EZ. stellulata.

200 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

It appears that the Hucalyptus coriacea consociation by no means possesses
the power of speedy renascence which characterizes Eucalyptus forests in other
regions. Whereas most Hucalyptus forests would have recovered from the fire
before the lapse of six years, here we found renascence to be tardy, and sometimes
not even to have started. Neither H. coriacea nor E. stellulata appear readily
to produce adventitious shoots from the stem, renascence taking place rather from
the root-stock; thus most of the regenerating H. coriacea forest showed young
shoots rising from the ground surrounding the old dead trunk, and only a few
feet high (Plate xviii, fig. 1).

In the more exposed areas no renascence was evident, and it seems that here
the severity of the climatic conditions has prevented it. In these areas the dead
trees are standing while the normal lower strata of the forest and especially the
Poa stratum have re-developed. It is likely that regeneration of the forest will be
slow, as it will now be chiefly dependent upon migration, followed by a precarious
ecesis (Plate xi, fig. 2).

The Eucalyptus stellulata consociation regenerates more rapidly, perhaps
because of the lower altitude and the consequently greater shelter of its environ-
ment; in this case a mass of shrub-like growth surrounded the base of the old
trunks, so dense as to cover the ground almost completely.

The regeneration of the lower strata was more advanced, and the burnt areas
did not appear to differ appreciably in these strata from those which were unburnt,
except that very dense shrub-societies had arisen in places. Where renascence of
the trees is slow dense societies of Veronica Derwentia, accompanied by Olearia —
megalophylla (f), Drimys (f), and Oxylobium ellipticum var. alpinum (0), are
generally characteristic features (Plate xiii, fig. 2). The rapid recovery of these
types is no doubt due to their resistant subterranean root-stocks or rhizomes.
The Eucalyptus Gunnii consociation is subject to more frequent burns and is more
akin in its reaction, to the Hucalyptus piperita Forest at Mount Wilson.

FACTORS CAUSING OR MODIFYING THE TIMBER-LINE.

In certain more exposed areas on the summits of the hills and on the higher
slopes, the tree-trunks in the Hucalyptus coriacea consociation appear to have been
killed by severe climatic conditions rather than by fire; for although regeneration
is taking place as usual from the base of the stem, there is no indication of fire
on the trunks, and moreover, these conditions are found only in exposed areas.
An analogous phenomenon is seen at altitudes of 6,500 feet, where the stunted
Eucalyptus forest has migrated during favourable seasons above the normal level
of the timber-line, and subsequently, possibly as the result of an unusually severe
season, has been entirely destroyed (Plate xvi, fig. 1).

These observations lead us to consider the factors which are effective in
controlling the tree-line in this region. Rhydberg (1914) has enumerated some
of those which he considered to be important in the case of the Rocky Mountains,
and it is of interest to note their significance in the case of the vegetation of the
Kosciusko Plateau. The chief of these factors described by Rhydberg are as
follows: (1) low temperature during the growing season; (2) shortness of the
growing season; (3) late frosts; (4) strong winds; (5) deep snow; (6) form of
precipitation; (7) large mountain masses; (8) physiographical barriers.

The timber-line on the Kosciusko Plateau varies from 6,100 feet in exposed
habitats to 6,500 feet in sheltered habitats. At these altitudes the forest thins out
and the trees are represented by low gnarled and stunted shrubby individuals;
the zone, however, between the absolute tree-line (where arboreal species disappear

BY J. MCLUCKIE AND A. H. K. PETRIE. 201

entirely) and the relative tree-line (where the arboreal species just cease to form
forest) is comparatively narrow. Exposure to desiccating westerly and north-
westerly winds, very cold in winter and hot in summer, determines the points
where the forest suddenly merges into isolated stunted individuals. In other |
alpine regions of the earth coniferous forests frequently exist above the angio-
spermic forest, but at Kosciusko the highest arboreal community is that composed
of Hucalyptus coriacea. :

Competition is of considerable importance in connection with the effect of the
various climatic factors on the tree-line; the transition zone is undoubtedly a scene
of constant struggle between the grassland and the scanty forest. In good seasons
seeds of Eucalyptus coriacea germinate and develop in the true grassland areas,
and if they survive, the grassland community becomes reduced to the status of a
subordinate stratum. No doubt this tree after difficult ecesis tends to carry the
forest upwards in this manner, but unfavourable seasons, when snow lies late in
summer as it occasionally does, and late frosts followed by a dry summer, cause
many of the trees to die down, perhaps for good, or perhaps till renascence sets
in; the grassland types then for the time resume their dominance, probably to be
subordinated again and again during long periods of slow oscillation. Throughout
the whole of this district, the higher forest tracts show abundant evidence of the
struggle against the grassland types. 3

The lowering of the temperature during the growing season (from November
to March), the first of Rhydberg’s factors, has an intermittent effect which probably
influences ecesis of the forest trees in their own or the grassland zone. The
shortness of the season, however, is more important in the Kosciusko district;
but both factors are closely related, for the prevalence of a low temperature during
the growing season as the result of late snow-falls or frosts, must inevitably curtail
the season for growth. Late snows are common in this area; in some years very
heavy falls take place in September, and lie long, water-logging the soil in certain
places and delaying the germination and growth not only of herbaceous but also
of arboreal types. In such seasons the small alpine and sub-alpine herbs are
several weeks later in flowering as compared with other seasons which have no
late snow-falls.

Strong wind in winter, especially during frosty periods, appears to be the
most important factor affecting the higher forest on the Snowy Mountains. Along
the upper limit of the forest, and where it passes into isolated groves or individuals,
the trees show the effect of wind by their gnarled, stunted, irregular, often prostrate
habit, and their frequently unilateral growth (Plate xix, fig. 2). This peculiar
and characteristic growth-form is probably due as much to the desiccating effect of
wind, by increasing transpiration from the youngest parts, as to its mechanical
action. This wind action prohibits the forest development above 6,100 feet on
exposed slopes and peaks, while on the leeward slopes, as has previously been
mentioned, the forest ascends to approximately 6,500 feet. The valley-heads and
erosion channels on the leeward side, however, are devoid of the forest which
occurs on the slopes, as has been described; this is no doubt due to the persistence
of the deep snow drifts in such positions, and the smothering of the root-systems
while the aerial organs are exposed to strong insolation and high transpiration.
This feature has been observed also by Rhydberg in the Rocky Mountains.

The significance of the amount of rainfall during the growing season is by no
means certain. On account of the high rainfall and low temperatures generally
associated with montane zones—to which the Snowy Mountains are no exception—
trees normally occurring in such habitats are usually found to have relatively high

202 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

moisture requirements and low temperature requirements (Pearson, 1920); and it
is noteworthy that the Hucalyptus coriacea forest maintains its optimum develop-
ment between about 4,500 and 5,600 feet, which is the zone of greatest rainfall.
Yet at the same time, although it may be that #H. coriacea has low temperature
requirements, and that this factor causes it to replace the H. Gunnii forest at

Baeckia Corsociation Callisemor Consociation

Epacris paludosa Consocies ~—____Epacris serpyllifolia Congocies

Richea Associes.
Sphagnum Stratum-Society
inthe Associes
Restio Associes WA
|
Hy polaena eee

Poa-Hypolaena Ecotone

Carex Consocies) Luzula Consocies

Poa Consociation

Text-figure 3. Illustrating the succession in the shallow valleys above 5,000 feet.
Baeckia should be spelt Baeckea.

BY J. MCLUCKIE AND A. H. K. PETRIE, 203

4,500 feet, its occurrence in exposed habitats near Goulburn, near Clarence, and
on the Monaro Plains, regions with a rainfall probably often bordering on 20
inches per annum during some months of the year, suggests that it is an
extreme xerophyte, more especially as the soil in these regions is dry and generally
of low moisture-retaining power. It is improbable that #. Gunnii or EH. stellulata
have lower moisture requirements, while our observations on H#. viminalis in other
localities seem to indicate that this has a moisture requirement somewhat higher
than most species of Eucalyptus. So far as the upper limit of the distribution of
Eucalyptus coriacea is concerned, however, it may be that the rainfall is of
importance, since it is to be expected that in the alpine region the effective rainfall
is low during the summer, the precipitation being mainly of the nature of light
showers which, while sufficing for the grassland vegetation, do not penetrate
deep enough to reach the roots of trees. .

THE MARSH VEGETATION.

In the flat, water-logged floors of the shallow valleys above an altitude of
5,000 feet, the Poa association, which occupies the lower portions of the slopes,
gives place to reed- and bush-swamp. This vegetation is exceedingly complex in
structure, comprising a large number of communities, all of which seem to be
in successional relation. The causes of this succession appear to be somewhat
obscure, and require detailed investigation. We have endeavoured in the present
paper, however, to characterize the communities and to indicate, to the extent of
our observations, the main lines along which succession seems to take place. No
doubt the amount, nature and distribution of the precipitation from year to year
and factors which modify the drainage system temporarily or permanently,
influence this vegetation materially.

The accompanying diagram (Text-fig. 3) will help to elucidate the remarks
which are made upon succession in the following detailed consideration of the
communities of this habitat.

Our present investigations indicate that there is little material physiognomic
or floristic variation in the communities of*the marsh vegetation throughout the
extent of their range, from 5,000 to about 6,500 feet, with the exception of the
Richea and Restio associes, which extend in a modified form to 7,000 feet.

THE POA-HYPOLAENA ECOTONE.
Where the Poa association of the lower parts of the slopes encroaches upon
the damp soil of the valley bottom it forms an ecotone in which it is mixed with
Hypolaena lateriflora, the dominant of the community which seems to be one of
the early stages in the development of the swamp flora. Hypolaena is a reed-like
plant about six inches in height, and grows densely between the tussocks of Poa.
Reference to the accompanying statement of the floristic composition shows
that a large number of the subordinates of the Poa consociation are absent from
the ecotone; these are apparently more suited to the drier soil, although in the
case of Stellaria pungens, which favours the damp erosion channels higher up the
slopes, it is possible that lack of drainage is an inhibiting factor. It will also be
observed that, in the case of a number of the components of the Poa consociation,
the frequency decreases on passing into the ecotone, which represents the limit of
their range; and in the same manner in this region first appear a number of the
subordinates of the Hypolaena associes, such as Oreomyrrhis andicola. Certain
types occurring in the Poa consociation, e.g. Prasophyllum fuscum, appear to find
a more favourable habitat in the ecotone, and their frequency increases. The
B

204 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

ny,

|

1

Pox

\thy

0

Pe
«j= Poa caespitosa tussocks. H Cry Comesperma retusum.
/ ey

H = ypolaecia laterifiora. €>- Epacris serpyllifolia.
Text-figure 4. Quadrat

Vo =" Lotss australis. ER- Craspedia Richet. 4 sq. metres of the Poa-
Hypolaena ecotone, near-

is est the marsh commun-
Lee Luzule campestris. 0 Olearia sp. ities. The spaces. be-
tween the tussocks of
T= nicnca Gunilt Q- Restio australis. Poa caespitosa are either
: bare, or occupied by
typically moisture-loving
D — Drosera peltata. Br- Brachycome sp. types, e.g. Hypolaena,
Drosera, Polytrichum,

ep LS paeh On a0. wat a eave, Richea and Liuzula.

Correct spelling should
Heo- Hemarthria compressa. S — _ S8tylidium lineare. be Craspedia Richea and

Baeckea.
Cs- Callistemon Sicberi. Wyk Sphagnum sp.

Eu- Buphrasia Brownii.

BY J. MCLUCKIE AND A. H. K. PETRIE. 205

tussocks of Poa are less continuous in the ecotone than in the Poa consociation,
the drainage channels between them which are occupied by Hypolaena and the
subordinates, being much wider (Text-figs. 4, 5).

2)

ae

& = Poa caespitosa tussocks.

Gg) Beccrie Gunviana, seedlings.

H = Ilypolaena laterifiora.

Eh) eruori paludosa, seedlings.

Ss = Stylidium lineare.

++

HE Polytrichune sp.

Mii Sphagnum sp.

iit i

hn

q
R
HM

+

He - Welichrysum sp.

Br - Brachycome

Eu - Huphrasia Brownii.

fic)- Ilemarthria compressa,

Text-figure 5. Quadrat of the Poa ecotone 1 m. square. The spaces between
the Poa tussocks are drainage channels. The presence of Restio, Hypolaena,
Epacris and Baeckea suggests the developmental nature of the community, while
in addition the mosses represent types not present in the Poa consociation. The
Restio associes (R) is developing somewhat precociously on the right at the
bottom.

Generally speaking this ecotone occupies the transitional habitat between that
of the Poa and that of the Hypolaena community. It may arise also, however, as
a result of a definite succession; local alterations in the drainage frequently lead
to increase in the soil-moisture-content of areas occupied by Poa, leading to
invasion by Hypolaena; as a consequence the same mixed community is found as
occurs in the ecotone, although here it possesses rather the character of a mictium
(Clements, 1916, p. 140).

THE HYPOLAENA ASsocIES. (Text-fig. 6.)

As the water-table reaches the surface of the soil, on passing from the
Poa-Hypolaena ecotone towards the centre of the marsh, the Poa disappears,
yielding the dominance to Hypolaena. This species forms a kind of uniform basal

206 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

stratum throughout all the communities of the marsh; and where it is not inter-
mingled with taller types it constitutes a distinct community. All stages of
abundance of these taller types may be found, especially in the case of Restio
australis; sometimes their distribution is so open that they rank merely as
subordinates in the Hypolaena‘associes although taller than the dominant, from
which condition they pass to being dominants themselves when in more close
formation.

Epacris serpyllifolia.

OO

Epacris serpyllifolia, seedlings.

4+4t

Callistemon Sieberi.

+
e+,

Callistemon Sieberi, seedlings.

ed
es

Epacris paludosa,

@#@

« +*

+
be
}

Epacris paludosa, seedlings.

Ae
a

Baeckia Gwiuniana.

er + tp

“4
+
+

¥4
*

Baeckia Gwuviana, seedlings.

Sphagnum sp.

S+@

Restio australis.

Poa caespitosa:;

o @&) »
|

Stylidium lineare.

Polytrichum Sp.

$f
t

O — Oreomyrrhis andicola.

Text-figure 6. Quadrat 1 m. square of the Hypolaena associes, showing
development of the Restio in the centre. A typical Sphagnum patch is seen on
the right, while the numerous Hpacris seedlings indicate that further development
is taking place.

Observations seem to indicate that Restio appears in flat areas where the
content of gravitational water is higher than it is normally in the Hypolaena
associes, and becomes more prolific with increased water supply. We suggest,
therefore, that the accumulation of water in parts, owing to the defective drainage
from the great quantities of melting snow on flat ground such as the valley
bottoms, causes Restio to spring up among the Hypolaena and to form a new
community.

Isolated societies of Sphagnum occur in the Hypolaena associes; in these
the normal subordinates of the associes, except Oreomyrrhis andicola, are often
less abundant, and occasionally Hypolaena alone remains. In some of the larger

BY J. MCLUCKIE AND A: H. K. PETRIE. 207

Sphagnum societies numerous seedlings of the epacrid Richea Gunnii appear, and,
since this type when found mature is usually growing in a dense cushion on old
Sphagnum beds, it is thought probable that the establishment of Sphagnum in the
Hypolaena associes paves the way for the development of a new community
dominated by Richea.

In somewhat drier areas than that in which the above succession takes place,
the Hypolaena associes contains numbers of seedlings, often exceedingly small, of
Epacris serpyllifolia or of HE. paludosa. There is no doubt but that this is evidence
of another succession taking place from Hypolaena to the Epacris associes; and
it is this evidence in particular, which clearly establishes the seral nature of
the Hypolaena associes. In the earlier stages of the developing Hpacris associes,
E. serpyllifolia seems often more abundant, with only occasional seedlings of
E. paludosa, although more rarely a pure developing consocies of the latter is
found; in the older stages, however, only the H#. paludosa consocies was seen,
although there is every reason to believe that the Hypolaena associes must be
capable of giving rise to both the Hpacris communities. It must be added at the
same time that there is undoubted evidence in places that the Epacris serpyllifolia
consocies gives place to the EH. paludosa consocies, the dominant of the later
community being a larger shrub; and it certainly seems in many cases that a
telescoping of this succession takes place in the developing Epacris associes, so that
the originally abundant seedlings of E. serpyllifolia subsequently give place to an
invasion of seedlings of H. paludosa, which is perhaps a more vigorous type. A
more probable explanation is that EH. serpyllifolia is better adapted to establish
itself first in the Hypolaena and that #. paludosa has to await further changes in
the conditions before it can colonize the habitat, after which it readily assumes
dominance on account of its larger size. /

It thus appears that the Hypolaena associes is a synthetic seral community,
which, collaterally with habitat changes in different directions, develops along
various lines.

THE RESTIO ASSOCIES.

Restio australis, which we have already referred to as dominating a community
developing in all probability from the Hypolaena associes, is a restiaceous plant
from one foot to eighteen inches high, growing densely and often occupying large
areas (Plate xii, fig. 2). The subordinates of the Restio associes will be seen from
the accompanying table to be much the same as in the Hypolaena associes, and
Hypolaena itself forms an almost constantly present lower stratum. FRestio,
however, occurs often with no subordinates as a pure society in little flat water-
holes so characteristic of the sub-alpine and montane marshy areas.

Sphagnum societies are occasional, in which communities most subordinates
except Hypolaena are absent; and judging by analogy with the previously
recorded observations in the Hypolaena associes, it is possible that these societies
may lead to the development of the Richea associes.

Seedlings of Epacris serpyllifolia and E. paludosa are abundant in places, so
that there appears reason to believe that when these have attained maturity the
Restio associes will have developed to a further stage in the sere dominated by
Epacris.

A large area of the Restio associes was observed where dead white gnarled
stems of Epacris indicated that this plant had previously made fair headway and
subsequently died out. Apparently its disappearance was followed by the estab-

208 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

lishment of the previous stage of the sere, viz., the Restio associes, suggesting
that a reversal of the changes in the habitat was the factor leading to the
destruction of the Epacris.

THE RICHEA ASSOCIES.

Richea Gunnii is a megaphyllous, unbranched or sparsely branched epacrid,
growing in dense clumps, usually on old Sphagnum beds (Plate xiii, fig. 1). The
habit of Sphagnum of forming mounds is well known, and Richea appears to find
its most favourable environment on these, sending long shoots upwards through
the Sphagnum; by this means the clumps of Richea reach a height of two or three
feet, although the normal height of the plant under ordinary conditions is usually
only about one foot. :

Richea grows so densely and in such a sodden substratum, that it ordinarily
has few subordinates. It seems, however, to consolidate the Sphagnum beds, and,
as the water in the soil gradually finds drainage channels by which to pass away,
other types are able to come in. The first to appear is Restio, after which comes
Hypolaena, suggestive of reversion of the sere owing to reversion of the habitat
changes. Hypolaena grows up through the Richea on long attenuated stalks, in
order to reach air and light, so that, although the plant is normally only about six
inches high this ecad attains a length of two or three feet. Where the drainage
and consolidation has proceeded further, Baeckea Gunniana gains entrance, as also
less frequently does Callistemon Sieberi. This seems to suggest a possibility of
Richea giving place to the Baeckea-Callistemon association, which is the climax
stage in the succession taking place in the swamp flora. Probably one may
explain in the same way the rather curious local presence among the Richea of
Ozylobium ellipticum var. alpinum, a type which usually inhabits much drier
places in the Hucalyptus forests.

Epacris paludosa also is present in some of the dying clumps of Richea, which
suggests succession from Richea to the Hpacris associes; this is rendered more
probable by the fact that Hpacris is usually an intermediate stage in the develop-
ment of the Baeckea-Callistemon association.

The distribution of Richea between 5,000 and 5,500 feet is much more restricted
than that ef the communities previously described, nor do the clumps often occupy
very extensive areas. From 5,500 to near 7,000 feet (the upward limit of its
range), however, the Richea associes gradually becomes more frequent and wide-
spread, until it eventually occupies extensive areas of the valley floors where the
habitat is sufficiently wet. Its structure in the alpine regions will be referred to
again subsequently.

THE EPACRIS ASSOCIES. (Text-fig. 7).

Reference has already been made to the presence of Hpacris seedlings in the
Poa-Hypolaena ecotone and in the Restio, Hypolaena and Richea associes,
which appear to develop into a community having Hpacris as dominant. So far,
the communities we have been considering have been distributed in zones of
increasing content of gravitational water in the soil; Hpacris, however, occurs in
drier soils more akin to that of the habitat of the Poa-Hypolaena ecotone and the
Hypolaena associes. The Hpacris associes may be a natural development of these
two communities in an unchanging habitat; and, indeed, every stage may be
observed between these two communities with only a few plants of Hpacris
scattered here and there (Plate xvii, fig. 1), to where it is abundant with only a

BY J. MCLUCKIE AND A. H. K. PETRIE. 209

subordinate stratum of either Poa with a little Hypolaena or Hypolaena with a
little Poa (Plate xv, fig. 2; Plate xvi, fig. 2). Consequently the transitional area
between the Poa consociation and the marsh may be occupied in places by a
Poa-Epacris ecotone rather than by a Poa-Hypolaena ecotone. The Epacris associes
also appears to follow the Restio and Richea associes coincident with the drainage
of the excess of gravitational water from the soil which takes place eventually as
the result of the gradual formation of erosion channels. In this case Poa is not
present, but later it is possible that Poa enters the habitat and becomes as abundant
as if the Hpacris had directly succeeded the Poa-Hypolaena ecotone.

(Ba) - Baeckia Gwiniana.

Be- Bacckia Guuniana, seedlings.

&5- Epacris serpyllifolia.

&- Epacris serpyllifolia, seedlings.

(Eh)- Epacris heteronema.

“EX Epacris heteronema, seedlings.

DI
4 (i) Wypolaena laterifiora.

R - Restiad australis.
Br- Brachycome sp.

C ~ Craspedia Richei.

D — Droscra peltatu.
Cs- Callistemon Sieber.
Ss — Stylidium lineare.
0 ~ Olearia sp.

Cr = Comesnerma rvetuswm,

Pou ccaespitosa,

Text-figure 7. Quadrat, 1 m. square of the Hpacris serpyllifolia consocies
in process of developing into the Baeckea consocies.

In certain drier parts of the swamp the Poa consociation was observed
merging into the ecotone, which in parts had apparently developed directly to
Restio with the omission of the pure Hypolaena stage, Poa still being present in
the Restio associes. In places the latter was developing into the Hpacris
serpyllifolia consocies, with scattered plants of the dominant between which Poa
was abundant. This seems to be a telescoping of the sere owing to drier edaphic
conditions.

Destruction of the Epacris paludosa consocies and its replacement by the Poa-
Hypolaena mictium was observed in several places, where the habitat was littered
with the dead gnarled and knotted stems (Plate xv, fig. 1). This destruction may

210 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

have been caused by snow lying long on the ground and killing out the Epacris.
The latter would then naturally be replaced by the Poa-Hypolaena community
which, as has been mentioned above, is akin in its habitat requirements.

As has already been mentioned, two consocies occur, dominated respectively
by HE. serpyllifolia and E. paludosa. The former seems to appear first and often
subsequently to give place to the latter, but in large areas EZ. paludosa is absent
and the EH. serpyllifolia consocies develops directly to the Baeckea-Callistemon
association, which usually follows the Epacris associes as the next and final stage
of the sere. Evidence of this interrelationship between these communities is seen
in the presence among the Epacris of locally abundant seedlings of both Baeckea
and Callistemon. Occasionally in the Epacris associes occur single mature plants

+

\

ha
\

\

SNC Sh: Gm MEAL:

\

,
\ -\
Pe Ng ¢
ee BS LY
eatt
fe + ++
t
(
} (
!

aN 4
W .
OGG
2 Fao
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g Peo
ss +
iN

ie}
(4
+

27 40 208
Q o o
acs LE. +

o a0
+ . 4 2O)
--\ 50% 5

oo eee

++ a Epacris serpyllifolia consociation.

4 - .Poa caespitosa consociation.
Op Poa-Hypolaena ecotone.

whe,
Aan. <~ Hypolaena and Restio associes.

C)- Richea associes.

632. Luzula consociation.
oo

ome

4 ‘ é
ome, ‘ contour line.
-o®

Text-figure 8. Diagrammatic Chart (not to scale) of a portion of the
marsh vegetation.

BY J. MCLUCKIE AND A. H. K. PETRIE. 211

of Callistemon Sieberi and Hakea microcarpa, shrubs about three feet high which
stand conspicuously above the dwarf EHpacris (Plate xvi, fig. 2).

The Epacris associes is one of the most widespread communities in the
marsh between 5,000 and 5,500 feet, and in many cases, indeed, may possibly be a
sub-climax. Above 5,500 feet, however, it gradually gives place to the extensive
Richea associes which apparently is more suited to the sub-alpine environment.
Text-figure 8 shows the interrelationship of the communities described in detail
in the preceding pages.

THE BAECKEA-CALLISTEMON ASSOCIATION.

This association appears to be the climax stage in the development of the
marsh vegetation, but rather curiously it is very restricted in its distribution, and
it is only here and there that conditions appear to permit the existence of the
dominants.

The association is present sometimes as a zone round the edge of the marsh
in a manner suggesting the occurrence of the Epacris associes, but only where the
ground has a well-marked slope; at other times it colonizes the margins of channels
of water running rapidly through the marsh. These facts of distribution suggest
that the association requires a more aerated soil than normally constitutes the
marsh; and this conclusion is supported by the similarity between the Callistemon
consociation of the association under discussion and the Callistemon society which
inhabits water-courses in the Hucalyptus forest higher up the slope (Plate xii,
fig. 1).

The dominants occur separately as consociations, the Baeckea consociation
being the more abundant and containing Callistemon Sieberi and Hakea microcarpa
as subordinates.

The number of species in this association is small, since the closeness of the
shrubs prevents the occurrence of lower strata and necessitates the display of all
foliage at one level; Richea, Hypolaena and Restio are present in the Baeckea
consociation growing up on long attenuated stalks through the mass of knotty
branches of the Baeckea, in the manner described for Hypolaena in the Richea
associes. The light ecad of Hypolaena in this community attains a height of
three feet, six times the normal height of the plant.

The destruction of Hpacris by severe conditions with consequent return to the
Poa-Hypolaena stage of the sere has already been described. A similar case was
observed of the destruction of the aerial shoots of Callistemon in a consociation of
that type, leaving the characteristic white stems standing erect throughout the
_ area. After the ground had been so bared, the Hypolaena associes had appeared,
and in parts had developed to the Restio associes, although not so far as. the
Epacris associes, H. serpyllifolia being only occasional. In the meantime a
fundamental difference between the epacrid and the myrtaceous type had asserted
itself. All our observations indicate that members of the Epacridaceae have
generally no means of renascence after destruction of the aerial shoots as is
characteristic of the Myrtaceae and many other families represented in Eucalyptus
forests; therefore, while regeneration of the Epacris associes has to await the
comparatively slow process of migration and ecesis, Callistemon, before even the
Restio stage had fully asserted itself, had commenced active renascence everywhere
by the production of adventitious shoots from the base of the stem, which no
doubt would rapidly grow up and restore the original structure of the community,
while the Restio and Hypolaena would be stamped out (Plate xiv, fig. 1). It is
interesting to note that Baeckea which was apparently occasional in the original

212 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

community, had survived destruction, suggesting that Baeckea is a more vigorous
type; a conclusion supported by its greater abundance at higher altitudes in the
Poa-Celmisia association.

THE LUZULA-CAREX ASSOCIES.

In flat shallow pools of water occur two closely related consocies dominated
respectively by the sedge-like Luzula campestris and Carex Gaudichaudiana.

The Luzula consocies occurs not only in the marsh vegetation, but also in little
flat hollows or natural drains in the Poa consociation or in flat areas of the
Callistemon Sieberi society in the Hucalyptus coriacea consociation.

The Luzula-Carex association does not appear to form a stage in the complex
sere already described, although it passes into it in several ways, as is about to be
mentioned. The floristic composition is also considerably different from that of
the various stages in that sere, since there are present a number of amphibious
or semi-aquatic types such as three species of Ranunculus, Utricularia, and even
the definitely hygrophytic Chara or Nitella.

In one example of the Luzula consocies Oreomyrrhis andicola was observed to
be equally abundant with Luzula. Since the leaves of that rosette-type form a lower
stratum than the Luzula we have termed it arbitrarily a sub-dominant; at the
same time when Oreomyrrhis is in flower or fruit, its pedicel is equally as tall as
the haulm of Luzula, so that during a large period of its existence it is co-
dominant. The community in which this observation was made was a less wet
habitat than is normally occupied by the Luzula consocies; and it is possible that
Oreomyrrhis has gained entrance to the community on this account. Increasing
drainage of the habitat in one part has resulted in the establishment of numerous
seedlings of Epacris serpyllifolia, indicating that the community is able to develop
into the #. serpyllifolia consocies and so link up with the main sere of the marsh
vegetation.

Another instance of this is exemplified by the presence of Restio in parts of
both the Carex and Luzula consocies, which of course on account of its larger size
soon becomes dominant, forming the typical Restio associes. Occasionally
Hypolaena is also locally abundant in the Carex consocies so that development to
the Hypolaena associes may possibly take place concomitant with consolidation of
the substratum and ablation of the surface water.

These various lines of possible succession are illustrated in Text-fig. 3.

Floristic Composition.
Reed Stratum. L. Gs

Restio australis R.Br. If

Epilobium glabellum Forst.
Grass Stratum.

ir}

Luzula campestris D.C. d f-e
Carez Gaudichaudiana Kunth. lf d
Hypolaena lateriflora Benth. — la
Rumex acetosella Linn. r —
Poa caespitosa Forst. If —_—
Plantago varia R.Br. ; If —
Brachycome sp. If —_
Oreomyrrhis andicola Endl. lsd —
Ground Stratum.
Ranunculus Millani_-F.v.M. a o-a
Utricularia dichotoma Labill. var. wniflora Benth. — la
Chara sp. or Nitella sp. — la
Velleya montana Hook. lf oo
Epacris serpyllifolia Labill. (seedlings) Ic —

Moss societies (chiefly Sphagnum and Polytrichum) ° —

213

Kk. PETRIE.

H.

BY J. MCLUCKIE AND A.

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THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

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BY J. MCLUCKIE AND A. H. K. PETRIE. 215

THE ALPINE VEGETATION.
THE POA-CELMISIA ASSOCIATION.

Above the tree-line in the neighbourhood of 6,000 to 6,500 feet, the open.
stunted Eucalyptus forest gives place to a vegetation composed mainly of low-
tussock grassland and herbs. This vegetation has a characteristically alpine
physiognomy, and forms a low dense covering over the treeless peaks which
reach an altitude of over 7,000 feet. Two consociations are found, the one being
dominated by Poa caespitosa, and forming a typical low-tussock grassland
community. This is in reality the same community as occurs below the Hucalyptus
forest on the slopes of the sub-alpine region, and as forms the ground stratum of
the forest itself. The composition of the subordinates, however, differs somewhat,
as will be seen by comparing the statement of the floristic composition at the end
of this section of the paper, with that of the ground stratum of the Hucalyptus
coriacea consociation and with that of the Poa consociation of the sub-alpine
region given with that of the marsh vegetation. The other consociation is
dominated by a creeping rosette composite, Celmisia longifolia, which forms a
dense felt-like growth. The Poa gives a rich olive-green to the hills in the
distance; Celmisia, however, with its glaucous leaves, thick white tomentum
and numerous inflorescences, appears in the distance as large bluish-white patches
scattered over the hillsides; this feature is illustrated in Plate xviii, figs. 2 and 3.

The subordinates of these two communities, as will be seen from the
accompanying statement of the floristic composition, are largely composed of the
same species, although it appears that the more delicate mesophytic types favour
rather the Celmisia consociation. The majority of the species conform to
characteristic alpine life-forms, as will also be seen from the statement of the
floristic composition. Apart from the tussock grass represented by Poa and the
mat grass represented by Hemarthria compressa, the dominant life-form is the
rosette herb, intermixed with a smaller proportion of cushion herbs. Shrubs are
only occasional and tend to be confined to the shelter of granite boulders; they
exhibit a marked degree of nanism, being either prostrate or in the form of low
cushions, usually occurring flattened close against the rock surfaces, in order
to avoid the effects of the strong winds (Plate xix, fig. 1). Among these shrubs
may be mentioned Grevillea australis, Podocarpus alpina, and the fern Polystichum
aculeatum.

The extreme shortness of the vegetative season at these high altitudes results
in every species flowering at the same period; consequently there is a wealth of
floral production unmatched in other regions of the State, more especially as the
flowers are mostly large and brightly coloured (Plate xix, fig. 3).

A certain amount of competition and overlapping takes place between the
Poa and Celmisia consociations, and often the two types are co-dominant. In
damper parts of the slopes and valley-bottoms in both consociations occurs the
dwarfed and prostrate HEpacris petrophila, which in parts becomes dominant, thus
forming a new consociation retaining the typical subordinates of the Poa and
Celmisia communities. The Richea Gunnii associes occurs also in the swampy
areas, where it is the main community, as has been mentioned earlier; this
species is also sometimes locally abundant in the Celmisia consociation. The
Epacris associes of the marsh vegetation, however, does not occur, and it appears
that the Richea associes forms the climax at these high altitudes. The Hypolaena
associes its found pure in one or two marshes, and this develops directly into the
Richea associes, which then contains Hypolaena in abundance. The EHpacris

216 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

petrophila consociation occurs associated with the Richea consociation in these
marshes but does not appear to be developmentally related. In water-courses
below the melting snow-drifts a Ranunculus rivularis society occurs with a
number of semi-aquatic subordinates, including Caltha introloba which is confined
to altitudes bordering on 7,000 feet.

On many drier areas Celmisia and Poa occur practically alone, since it appears
that the subordinates are the less hardy and more mesophytic types.

During nine months of the year the alpine region is covered in snow which
in places frequently lies as drifts throughout the summer. The effect of this on
the vegetation is naturally very pronounced. All the herbaceous types die down
when the snow comes, but perennate from season to season by subterranean organs
of propagation. Thus an area from which the snow has just been melted is seen
to be covered with the withered remains of vegetation; and later appear the young
shoots of Poa, etc. In some areas where snow has remained continuously for
several years these withered remains have very largely disappeared, and no doubt
the perennating organs are here entirely killed out, so that a new migration will
have to take place. These occasional patches are the only areas of granite soil
which, in summer-time, are not clothed with a dense carpet of vegetation. Contrary
to this feature of the lifehistory of the herbaceous plants, the shrubs are able to
withstand the effects of the snow covering, without even losing their leaves, since
no periodically deciduous species occur in the New South Wales flora: At the
same time the shrubs are often killed during severe seasons, as is evinced by
the bleached, gnarled and thick woody stems which are often to be seen.

The Poa and Celmisia consociations occur also on a large area of slate in the
alpine region, where they possess practically the same features. A number of
smaller differences in floristic composition appear to exist, as will be seen from
the accompanying lists; but we have no definite record of any plant being confined
to the consociation on the slate, nor have we definite record of any species not
occurring on the slate. It is probable, however, that further work may modify
this statement, since our observations on the composition of the slate flora are
incomplete.

In the alpine region numerous very typical structural and growth features are
exhibited by the herbaceous and shrubby plants. The habit of the shrubs varies
with exposure to wind. In the most exposed situations where the desiccating effect
of wind is predominant and retards normal elongation of the aerial shoots, nanism,
or stunted, dwarfed and distorted growth results. The larger shrubs such as
Phebalium ovatifolium, Orites lancifolia, Prostanthera cuneata, Podocarpus alpina
and Grevillea australis assume a close, dense, caespitose habit, which favours
reduction of transpiration and natural protection from wind-action. Forms like
Pimelea alpina, Lissanthe montana, and Kunzea Muelleri frequently assume a
typical espalier habit, prostrate and straggling, and raising their growing tips
only a few inches above the level of the soil or protective rock.

Several mat-forming herbs occur in the alpine zone, being generally very
dwarfed forms which cling closely to the surface of the soil. The most important
of the plants in this category are Stackhousia pulvinaris, Raoulia catipes, and
Epilobium confertifolium.

The rosette leaf habit is represented in a number of types, as will be seen
from the accompanying statement of the floristic composition. The majority
of these plants are characterized by deep tap roots.

Sclerophylly is a characteristic feature of the shrubs, while many types, both
shrubby and herbaceous, have extensive and dense coverings of silky or woolly

Species.

Acaena sangwisorbae Vahl.
Aciphylla glacialis F.v.M.
Astelia alpina R.Br. ......
Brachycome cardiocarpa
F.v.M. var. alpina
Brachycome scapiformis D.C.
Barbarea vulgaris R.Br.
Carex breviculmis R.Br. ...
Celmisia longifolia Cass.
Claytonia australasica Hook.
Craspedia Richea Cass.
Craspedia Richea Cass. var.
alpina Benth.
Drapetes Tasmanica Hook.
Epacris petrophila Hook. ..
Huphrasia Brownii, F.v.M.°
Galium wumbrosum Sol.

Grevillea australis R.Br.

Helipterum incanum D.C...
Hemarthria compressa R.Br.
Hymenanthera dentata R.Br.
Kunzea Muelleri Benth.
Lissanthe montana R.Br. ..
Lycoperdon sp.
Lycopodium clavatum R.Br.
Myosotis australis R.Br. ...
Olearia floribunda Benth. ..
Oreomyrrhis andicola Endl.
Oreomyrrhis pulvinifica F.v.M.
Orites lancifolia F.v.M.
Pimelea alpina F.v.M.
Pimelea axiflora F.v.M. var.
alpina F.v.M.
Phebalium ovatifolium F.v.M.
Poa caespitosa Forst.
Podocarpus alpina R.Br.
Polystichum aculeatum (L.)
Schott.
Polytrichum sp.
Pentachondra pumila R.Br.
Prostanthera cuneata Benth.
Ranunculus anemoneus F.v.M.
Ranunculus dissectifolius
F.v.M.
Ranunculus Gurnnianus Hook.
Ranunculus Muelleri Benth.
Azorella dichopetala Benth.
Ranunculus sp.
Raoulia catipes Hook.
Richea Gunnii Hook.
Senecio pectinatus D.C.
Stackhousia pulvinaris F.v.M.
Taraxacum dens-leonis Desf.
Viola betonicifolia Sm.
Danthonia robusta F.v.M...
Epilobium confertifolium
Coprosma Nertera F.v.M...
Lycopodium Selago, Linn...

—_—_——————————————

BY J. MCLUCKIE AND A. H. K. PETRIE.

Floristic Composition.
Poa-Celmisia Association.

Life-form.

Creeping mat herb
Rosette herb
Rosette herb

Rosette herb
Rosette herb .....
Low erect herb

Sedeere io cee
Creeping rosette herb
Mat herb

se ee ee

Rosette herb ......
Prostrate shrub
Prostrate shrub
Low erect herb ....
Low herb .........
Caespitose or pros-
trate shrub .....
Creeping rosette herb
IOWA LASS eee
Espalier shrub
Hspalier shrub
Hspalier shrub
italy wary eicueencee:
Decumbent herb
Herb
Espalier shrub
Rosette herb
Rosette herb
Caespitose shrub
Espalier shrub

Espalier shrub
Caespitose shrub
Low-tussock grass
Caespitose shrub

ee ew ew we

ey

Hspalier shrub
Caespitose shrub
Rosette herb ......
Rosette herb
Creeping rosette herb
Rosette herb
Rosette herb
Rosette herb
Mat herb
Low caespitose shrub
Erect herb
Mat herb
Rosette herb
Creeping rosette herb
Tufted grass
Mat herb 5.250...
Espalier shrub

see eee

|
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Poa | Celmisia
consociation.| consociation.
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la

217

Slate.

Celmisia

218 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

hairs on their inflorescences, leaves and young shoots. Examples of this class are
Celmisia longifolia, Helipterum incanum, Podolepis longipedata, Craspedia Richea
var. alpina, Raoulia catipes, Oreomyrrhis andicola and Ranunculus Muelleri. The
relation of hairiness of alpine and sub-alpine plants to protection against intense
insolation, and consequent excessive transpiration and inhibition of growth, is so
well known that it needs no discussion here.

Epacris petrophila Consociation.

Epacris petrophila Hook. d

Celmisia longifolia Cass. f-e
Richea Gunnii Hook. la
Craspedia Richea Cass. o-f

Richea Associes.

Richea Gunnii Hook. d
Restio australis R.Br. la
Sphagnum sp. (9)

Ranunculus rivularis Society.

Herb Stratum.

Ranunculus Gunirianus Hook.

Brachycome cardiocarpa F.v.M. var. alpina f
Ground Stratum.

Ranunculus rivularis Banks and Sol.

Caltha introloba F.v.M.

Drosera Arcturi Hook.

Sphagnum sp.

oov em

THE EPACRIS PETROPHILA CONSOCIATION.

The summits of certain of the peaks in the slate region of the Plateau
represent an extremely barren habitat: the ground is composed of the upturned
edges of the strata of slate, and there is practically no soil. The conditions seem
too severe for the grassland which tends to accumulate the raw humus of a
moisture retaining soil, and a few of the hardy species form an open community
which has the characteristic physiognomy of fell-field or alpine rocky desert
(Plate xvii, fig. 2). None of the individuals are more than three or four inches
in height, and they are scattered intermittently over the rocky ground. The
available moisture is less than elsewhere, hence the plants are mostly xerophytes,
especially the dominant, Epacris petrophila. It is curious in this connection that
this species should have been characteristic of the damper slopes in the Poa-
Celmisia association.

Although the majority of the components of the community were found in
the Poa-Celmisia association, the great difference in physiognomy marks it out
as a distinct association, distinct even from the consociation of the Poa-Celmisia
association having the same dominant.

A remarkable feature is the sudden replacement of the Hpacris petrophila
association by the Poa consociation on passing to a more sheltered habitat over
the brow of the hill, the ecotone region being only a few yards in breadth.

Floristic Composition.
Epacris petrophila Hook.
Colobanthus subulatus Hook.
Carez breviculmis R.Br.
Azorella dichopetala Benth.
Azorella cuneifolia F.v.M.
Poa caespitosa Forst.
Brachycome sp.
Buphrasia Brownii F.v.M. very small form
Lichens

lay bei (a) (a) (co) (o) Ley ia-y jen

BY J. MCLUCKIE AND A. H. K. PETRIE. 219

Summary.

The paper comprises an account of the plant communities of the Kosciusko
Plateau. ;

Three regions are recognized on the plateau, viz., the alpine, sub-alpine and
montane zones.

The montane zone and the slopes of the ridges in the sub-alpine zone are
occupied by Eucalyptus forests, the former mainly by a Hucalyptus Gunnii
consociation, the latter by a Hucalyptus coriacea consociation.

Accounts are given of the structure and adaptations of these forests, including
certain features of regeneration, and of the factors controlling the timber-line at
the upper limit of the sub-alpine zone.

The flat bottoms of the shallow valleys of the alpine and sub-alpine regions are
occupied by a: marsh vegetation comprising reed-swamp and dwarf-shrub heath.
The interrelationships of a number of seral communities in this habitat are
discussed at some length.

The alpine zone, extending from the upper limit of arboreal vegetation to
the summit of the plateau, is occupied mainly by a Poa-Celmisia association. The
‘Poa consociation forms a low-tussock grassland community, and occurs also in the
sub-alpine zone between the Hucalyptus forest on the slopes and the marsh
vegetation in the valley bottoms; the Celmisia consociation forms a mat-herb
community and is confined to the alpine region.

Acknowledgements.

The authors desire to record their thanks to Professor Griffith Taylor and his
associates for the privilege of reproducing their reconnaissance map and the block
diagram of the Kosciusko Plateau, and to Mr. EK. Cheel of the National Herbarium
for kindly identifying a few of the specimens.

References.

CLEMENTS, F. E., 1916.—Plant Succession. Carneg. Inst. Wash. Publ. 242.

HeumMs, R., 1896 and 1897.—The Australian Alps or Snowy Mountains. Jour. Roy. Geog.
Soc. N.S.W., Vol. VI, 4, 1896, and 5, 1897.

MAIDEN, J. H., 1898-99.—The Flora of Mt. Kosciusko. Agric. Gaz., N.S.W., 1898 and 1899.

PEARSON, 1920.—Factors controlling the Distribution of Forest Types. Hcology, 1.

RHYDBHRG, 1914.—Review of two papers in Journ. Ecol., II, p. 140.

TAYLOR, G., BROWNE, W. R., and JARDINE, F., 1925.—The Kosciusko Plateau, a topographic
reconnaissance. Proc. Roy. Soc. N.S.W., 59, p. 200.

EXPLANATION OF PLATES X-XIX.
Plate x.

Map of Kosciusko Plateau (20 miles by 6 miles), where the elevations are represented
by form-lines or approximate contours.

Plate xi.

1. Slope of a shallow valley at about 5,100 feet. The Hucalyptus coriacea
consociation, with its characteristic shrubs, is seen in the rear. The typical life-form
of the dominant is well illustrated in the comparatively large tree on the left. The
Poa consociation is present on the slopes on the right and left, while the marsh
vegetation occupies the level ground where water collects. The low dark heath-like
portion is the Hpacris association, while the Hypolaena associes and such communities
constitute the remaining portion of the marsh vegetation.

2. A shallow valley at about 5,500 feet, showing destruction of Hucalyptus coriacea
by severe climatic conditions. The lower stature of the trees at this altitude compared
with those of the previous photograph is evident. The replacement of the forest by the
Poa consociation on the lower parts of the slopes is shown; the marsh vegetation is

Cc

220 THE VEGETATION OF THE KOSCIUSKO PLATEAU, i,

sparingly developed, as the drainage is comparatively good in this instance, but darker
patches of it are seen in the middle distance. The occurrence of isolated trees on small
raised drier hummocks in the Poa consociation is a conspicuous feature.

Plate xii.

1. View at 5,200 feet, showing the Eucalyptus coriaecea consociation in the back-
ground, and the Callistemon consociation in the foreground.

2. The Restio associes in gently undulating country at 5,500 feet. The Poa and
Eucalyptus coriacea consociations are seen on the higher ground in the distance.

8. A shallow valley at 5,600 feet showing the Hucalyptus coriacea forest on the
ridge and the Poa consociation in the foreground. The lower stature of the trees than
in Plate xi, fig. 1, is again evident. The tussock-like nature of Poa caespitosa is more
noticeable at this higher altitude on account of the greater abundance of drainage water.
This is particularly seen in the foreground.

Plate xiii.

1. A granite boulder in a marsh at 6,000 feet, showing Baeckea Gunniana flattened
against it, in order to obtain protection from wind. Richea Gunnii is seen in the right
foreground.

2. A society of Veronica Derwentia in a portion of the Hucalyptus coriacea
consociation at 5,500 feet, which has been killed by fire. Poa is seen to occupy the
ground between the plants of Veronica.

Plate xiv.

1. A regenerating plant of Callistemon Sieberi, whose aerial portion has previously
been killed by severe climatic conditions.

2. The Snowy River at 4,000 feet, showing the Hucalyptus Gunnii consociation
clothing the slope in the background, and the Leptospermum society lining the river
bank. The difference in the topography from the higher altitude, as well as the denser
forest is evident. The bare nature of the forest floor in the H. Gunnii consociation is
seen on the right.

Plate xv.

1. In the middle distance is seen the Epacris paludosa consocies in flower. The
foreground was originally occupied by this type, which has, however, died out, as is
evident by the dead branches, and has been replaced by the Poa-Hypolaena mictium. The
Eucalyptus coriacea is well developed in the background. Altitude 5,100 feet.

2. Another view of the Hpacris paludosa consocies in flower, passing through the
Poa-Hypolaena mictium which reaches as far as the foot of the white tree on the right
and then merges into the Poa stratum society of the forest.

Plate xvi.

1. A view in the alpine region at 6,700 feet. The white patches on the steep slope
on the left are dead trunks of a low forest of Hucalyptus coriacea, which has migrated
upwards and subsequently been killed by severe seasons. In the foreground is seen the
Celmisia consociation, the dominant being admixed with Poa. The distant ranges are in
Victoria.

2. The Epacris paludosa consocies, 5,100 feet. A bush of Hakea microcarpa is seen
in the background. a

3. A view in the Hucalyptus Gunnii consociation, 4,000 feet, showing the tall trees
and the ground almost devoid of vegetation.

4. A shallow valley at 5,800 feet. The Hucalyptus coriacea consociation is seen to be
here very stunted and confined to the ridges between the erosion-channels running down
the slope. Between the patches of forest are dense societies of Prostanthera cuneata
which extend down over the Poa consociation on the lower part of the slope, On the
floor of the valley in the creek bed occur the Restio and Hypolaena associes.

Plate xvii.

1. The Poa-Hypolaena ecotone developing into the Hpacris associes. Plants of
Epacris paludosa are scattered through the Poa. <A plant of Callistemon Sieberi is seen
in the foreground.

BY J. MCLUCKIE AND A. H. K. PETRIE. 221

2. Summit of a ridge in the slate area at the summit of the Plateau, occupied by
the Epacris petrophila consociation. The area is seen to have the characteristic
physiognomy of fell-field, the plants being scattered openly among the projecting edges

of the folded slate strata.

Plate xviii.

1. A portion of the Eucalyptus coriacea consociation at 5,200 feet, in which the trunks
of the trees have been killed by fire. Regeneration is seen to be taking place from the
rootstocks.

2. The Poa-Celmisia association above 7,000 feet. The Celmisia, in flower, is seen
to alternate with the much more sombre-coloured Poa consociation. The characteristic
patchwork effect of the alternation of these two communities is seen on the distant hills.

38. A view of the alpine region from Mount Kosciusko, 7,328 feet. The patchwork
effect of the Poa and Celmisia consociations is again seen covering the distant hills.

Plate xix.

1. Podocarpus alpina in the shelter of a granite boulder, 7,000 feet. Under the rock
on the right is the habitat of Polystichum aculeatum, the only fern occurring in the
alpine region. ,

2. Scene at 6,000 feet showing a few isolated, gnarled and stunted trees of
Eucalyptus coriacea in the foreground, surrounded by a dense society of Prostanthera
cuneata mixed with Phebalium ovatifolium. Forest clothes the lower ridges on the left,
but the others are practically above the tree-line. Water is seen lying in a marsh on
the flat ground to the left.

3. The Celmisia consociation at 7,000 feet, with dense societies of Huphrasia Brownii,
all the plants being in flower.

[The following names on the text-figures, submitted incorrectly, were not noticed
until the blocks had been made. Text-fig. 2, Stellaria; Text-fig. 3, Baeckea; Text-fig. 4,
Baeckea and Craspedia Richea; Text-fig. 5, Baeckea and R = Restio australis; Text-fig. 6,
Baeckea; Text-fig. 7, Baeckea and Craspedia Richea.—b. ]

AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES. No. v.
By H. J. Carter, B.A., F.E.S.
(Six Text-figures. )
[Read 27th July, 1927.]

BUPRESTIDAH.

Mons. Théry has lately published (Ann. et Bull. Soc. Hnt. Belg., 1926) a
notable contribution to our knowledge of the Chalcophorini, that includes the
useful, but sometimes provocative, task of research on synonyms. With some
diffidence I suggest the following notes on the Australian species of Cyphogastra
known to me.

According to Théry both @. venerea Thoms. and C. macfarlani Waterh. are a

subspecies of C. farinosa F., macfarlani being considered as a variety of Thomson’s
species. I find the limit of the term subspecies difficult to define, but the term is
a convenient one as usually applied to a clear geographical race, that the student
of evolution believes to show the origin of a distinct species. With this under-
standing both venerea and macfarlani seem to me to deserve equal rank, at
least as subspecies. Macfarlani, besides having the extra lateral pubescent
impression (entirely wanting in venerea) differs (at least in my specimens) in
the outline of the upper surface as viewed sideways; venerea being reflexed at
apex, while the outline of macfarlani appears flat, or even deflexed at apex.
; C. vulnerata Théry.—Théry now considers this as a subspecies of C. pistor
L. and G., but I would claim specific rank for this species, that is, if I correctly
identified it from description when I figured it (These ProckEpines, 1921, p. 305)
alongside C. browni Cart. Thus from C. pistor it differs as follows:

A. By markings—lateral sulcus continuous from near shoulder to near apex;
two basal marks elongate instead of round; two premedial marks, linear in
vulnerata, oval in pistor. B. Apices of elytra much more strongly spinose. C. The
outline (as viewed sideways) convex, the highest point at basal third, the outline
descending lightly to front, more strongly to apex (in pistor horizontal with a
slight preapical concavity). D. Form of prothorax—nearly straight in vulnerata,
more arcuate in pistor. Of the above C appears of the greatest value.

C. browni Cart.—I consider this at least of sub-specific value, the several
examples captured by Mr. Brown showing the constant differences from pistor
that were stated in my description. I would reserve the term variety for those
examples that show more superficial differences, chiefly of colour and size, so well
known in the genus Stigmodera, and are generally without any zoo-geographical
significance.

Cisseis viridipurpurea Cart.—I took a number of this species on the Geraldton
wax plants (Chamaclaucium uncinatum) at Geraldton in September, 1926, the

described type was a J, in the ? specimens (not described) the head and pronotum
are concolorous with the elytra.

BY H. J. CARTER. 223

CHRYSOBOTHRIS.

In my revision of this genus (These PRocEEDINGS, 1925, p. 227) I quoted Mr.
J. Clark as to their habit of frequenting Acacias. Mr. T. G. Sloane has lately
supplied me with the following field notes on this subject that are characteristic
of this accurate observer, in reference to specimens of C. mastersi Macl. taken
recently at Moorilla, near Young, N.S.W.

“T took over a dozen during a week in February in blazing hot weather. I
got all the specimens in the bright sunshine from 11 a.m. to 2 p.m. on a large
fallen limb of Hucalyptus albens. The Chrysobothris was running on the rough
bark of the fallen branch (had nothing to do with the leaves) and I believe lays
its eggs in erevices of the bark. The Anilara (A. anthaxoides Théry) was in the
same place on the same limb and with the same habits”.

The following notes on what is probably the’ same Chrysobothris were made
at Mulwala (Murray River):

(1) December 17, 1880.—‘‘Saw the first 6-spot Buprestis of the season on a
fallen bough (Eucalyptus meliodora)”’.

(2) December 26, 1880.—‘‘Captured 4 specimens of the 6-spot Buprestis on a
fallen limb of a yellow boxtree (H. meliodora). A very hot day, wind from
N.W.; thermometer 106° in the verandah”.

(3) January 23, 1887.—“A bush fire has been burning since the 19th; at some
green trees (H. meliodora or E. hemiphloia) that were burned down I took 3
dozen specimens of the 6-spot Buprestis; also 5 dozen specimens of Merimna atrata.
One I watched caught and devoured a small buprest (Anilara sp.)”.

Mr. Sloane further notes in his letter, ‘‘wattles were plentiful at Mulwala and
I never saw the local Chrysobothris on or about them”.

SrigMoDERA (THEMOGNATHA) DUCALIS, n. sp. (Text-figure 1.)

¢. Elongate oblong ovate, head and pronotum nitid black (the latter with
bluish gleams at sides), elytra orange red, with narrow basal margin, the suture
narrowly, a large oval or oblong spot behind scutellum, sometimes with an
extension to the scutellum, two oblique linear humeral marks, a wide postmedian
fascia extending over about three-quarters of the width, black or blue black, the
apex narrowly blue black; underside and legs mostly dark bronze or greenish
bronze, the apical segments of abdomen more or less red.

read closely, regularly, not coarsely, punctate, front lightly channelled between
the eyes.

Prothorax (13 x 9-5 mm.), lightly convex, apex and base bisinuate, sides
moderately and regularly rounded, with narrow coarsely punctate lateral
explanation, all angles acutely produced, disc with fine shallow punctures, becoming
coarse and rugose at sides, basal half with a finely cut, shallow, medial sulcus.
Scutellum transversely oval and convex, nitid and impunctate. Hlytra of same
width as prothorax at base, soon widening behind shoulder, lightly compressed
at middle and feebly widening on posterior third, apices each bidentate (the
sutural tooth the longer) with a wide truncature between the teeth, elytra slightly
dehiscent at apex, striate, each with ten striae besides a short scutellary stria, the
fifth interval irregularly rugose-punctate, lateral intervals each with a single row
of fine punctures, the medial intervals scarcely, or very sparsely, punctate.
Underside closely and strongly punctate, the sternal regions rather densely clothed
with long silvery hair; abdomen glabrous and less strongly punctate; apical
segment arcuately excised.

224 AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES, V,

2. In the two examples examined, the underside is entirely dark bronze,
except for a little red near the lateral edges of the three apical segments, while
the abdomen is extended considerably beyond the elytral apex, with a protruding
Ovipositor. In one’ example the oblique humeral marks form a narrow lunate
fascia, connected with the hind edge of the post-scutellary patch. Dimensions:
6, 27-40 x 10-5-15 mm.; 9, 44-48 x 15-17 mm.

Habitat—wWestern Australia, Zanthus (Transcontinental Railway).

Five examples examined, the first of which was sent for description by Mr.
F. E. Wilson. It cannot, I think, be confused with recorded species, and should be
placed near S. spencei L. and G. in my tabulation. The form, sculpture and nitid
surface of the pronotum is somewhat as in S. oleata Blkb., but it is less widely
rounded, less convex, with a more explanate lateral margin. Holotype ¢ in Coll.
Wilson. Allotype 9 and paratype ¢ in National Museum, Melbourne. Paratypes
dg and 9 in Coll. Carter.

Text-figure 1. Stigmodera ducalis, n. sp.
Text-figure 2. Stigmodera miranda, Nn. sp.
Drawings by C. Deane.

STIGMODERA (THEMOGNATHA) MIRANDA, n. Sp. (Text-figure 2.)

dg. Oblong, lightly attenuated in front and behind, very nitid; head, pronotum
and basal three-quarters of elytra dark blue, subapical fourth and margins (widely
extending to a humero-basal spot) sanguineous, suture and extreme apex narrowly
blue; beneath rather brilliant bronze, sometimes with greenish tinge, flanks of
prosternum cyaneous, apical segments of abdomen more or less sanguineous; sternal
area (especially prosternum) rather densely clothed with long, upright, silvery
hair.

Head, having a light smooth impressed line between eyes, front feebly
concave, moderately punctate and rather strongly pubescent. Prothorax wide and
subexplanate, lightly convex, apex and base bisinuate, anterior angles obtuse,
posterior subrectangular, widest behind middle, hinder part of sides parallel,
thence arcuately narrowed to apex, disc with finer punctures than with its allies,
these interspersed with fine vermiculate smooth spaces; punctures and vermicula-
tions becoming stronger towards lateral margins; a smooth impressed medial
line on basal half. Scutellum subcircular, nitid and impunctate. Elytra as wide
as prothorax at base, soon widening and strongly sinuate before middle, rather
widely rounded behind, each apex widely truncate between two short teeth,

BY H. J. CARTER. 225

margins entire; striate, intervals strongly convex, the sixth broken up into
irregular ridges and punctures as also the basal parts of the second and fourth;
intervals (chiefly the lateral) with a few small punctures. Underside: The pro-
and meta-sternal punctures largely concealed by hair, but evidently not coarse,
abdomen densely covered with moderate sized punctures and a little strigose;
apical segment arcuately excised.

© differs in its larger size, less amount of red on apical segments (here
only showing at sides), apical segment of abdomen elongate and rounded behind
(with projecting sternite), more brilliantly nitid and much less hirsute beneath.
Dimensions: 3g, 40-42 x 15-16 mm.; 9, 46-50 x 17-19 mm.

Habitat—Western Australia; Zanthus (National Museum presented by Mr.
J. Lambe and Coll. Carter from R. Halse). Five examples before me, 3 64,
2 29, of which four are from the National Museum. Holotype ¢ and allotype ? in
the National Museum, Melbourne. I am indebted to Mr. Cedric Deane, C.E., of
Melbourne, for the excellent figures of S. ducalis and S. miranda. A magnificent
species that can be distinguished from its allies by the following table.

Stigmodera (Themognatha), colour largely black, or very dark blue, with
sanguineous markings.

1. Sides of elytra almost straight, margins of pronotum in general (in five out of six
examples! before me)) narrowilye reas seme sche eicies =: cleieclce elie rectipennis Blkb.

Sides of elytra clearly sinuate, pronotum without red margins .................. 2

2. Apices simply rounded, intervals regular throughout, colour largely black ..........
61850. DIONE CLARO O OIE OSG ROR EREHO EEO 1010.00 6-0 DOIN GiD1G-0'6 60'S Ch ORCI UEC CRAIC E Oicro ined princeps Blkb.

Apices not simple, sixth interval (besides others at base of elytra) irregular, colour

DUC crore ite Be cc ch aie iia Aaa eh athena Monee Mec tine te mrearem ete eietrcttatl amc suse aciva pis cigs tevenepecen siren suc uet ra Versi Wetag car 3

3. Pronotum strongly punctate (rugose at sides) with a faint smooth medial line,
UNAerside? CY ANEOUSH azo js sieusietsee he oat dabe eke ober ai oh caeree rasta eels) ules *bonvouloiri Saund.
Pronotum finely punctate, with impressed medial channel on basal half, underside
ACNECOUS aA DICAliMSESIMENES Tre Cerne emer adi eel cieic icicle euetcuer etenel el chake miranda, n. sp.

*JIn bonvouloiri the pattern of the upper surface is variable, from that portrayed
in Saunders’s figure to examples in which the elytra are mostly blue with a greenish
tinge, the apex only red.

STIGMODERA (CASTIARINA) DEUQUETI, n. sp. (Text-figure 3.)

Oblong, moderately attenuated in front and behind; head, pronotum and
scutellum golden bronze, underside golden green, antennae and legs green, tarsi
blue, elytra yellow with the following markings bright green, basal border and
extreme apex (both narrowly), an elongate, 3-pronged fork-like mark, the outside
prongs forming elongate vittae terminating at the humeral callus, the middle
prong covering the suture to the base, a sagittate subapical mark connected
narrowly at suture with the base of the fork.

Head excavate and canaliculate, moderately punctate. Prothorax widest near
base, thence subparallel to base and arcuately narrowing to apex, lightly convex,
apex feebly, base more strongly bisinuate, anterior angles subacute, posterior
subrectangular, disc without defined medial line (except a short, smooth area),
a medio-basal and two large latero-basal foveae; surface punctures defined and
distant on basal half, closer and finer towards apex, strong and close at sides.
Scutellum triangular, almost impunctate. Hlytra of same width as prothorax at
base, widening at the humeral swelling, slightly compressed behind this, apices
finely bispinose each with a short, oblique lunation, subapical margins entire;
striate-punctate, the seriate punctures close, intervals impunctate, the third and
fifth roundly convex, the rest flat except at apex. Underside very lightly pubescent,
finely and evenly punctate. Dimensions: 12 x 5 mm.

226 AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES, V,

Habitat—New South Wales, Armidale (Mr. C. F. Deuquet).

I name this brilliant species after its discoverer, who has generously given
a pair to me and the holotype ¢ to the Australian Museum. The elytral pattern
is constant (Mr. Deuquet has several other examples), and is near that of the
tricolor Kirby group, though not quite like any species known to me. The ground
colour is that of confusa Waterh., in which the pronotum is described as “globose”
and the subapical margins serrate, also with a different elytral pattern. This last
must be very near that of S.’clancula Obenb., but in this species the head,
prothorax and elytral pattern are blackish (‘“‘schwdrzlich’), the underside is
bronze, little shining (‘‘wenig gldnzend’’), the elytra appears to differ in form
from deuqueti, each with a single tooth at apex, and the habitat is given as
Queensland. Holotype ¢ in the Australian Museum.

Text-figure 3. Stigmodera dewqueti, n. sp.
Text-figure 4. Stigmodera fossoria, n. sp.
Text-figure 5. Stigmodera dilatata, n. sp.
Text-figure 6. Stigmodera flavo-viridis, n. sp.

STIGMODERA (CASTIARINA) FOSSORIA, nh. Sp. (Text-figure 4.)

Narrowly oblong; head, pronotum, scutellum, elytral markings, underside
and appendages dark green; elytra testaceous, with a wide, sagittate subapical
mark, the apical three-fourths of the suture, and the extreme apical margin (very
narrowly) dark green, the latero-apical margins slightly tinged with red.

Head deeply channelled and excavate, more brightly metallic than pronotum.
Prothoraz rather strongly convex, apex sub-truncate, base strongly bisinuate,
widest at middle, sides well rounded, all angles, seen from above, obtuse; disc
uniformly and closely punctate, medial sulcus distinct on basal half (traceable
throughout in one example). Elytra as wide as prothorax at base, soon widening,
the sides nearly straight for the greater part, feebly compressed behind shoulders,
striate-punctate, intervals minutely punctate, the short scutellary interval, the
third, fifth and seventh more strongly convex, wider and less punctate than the
alternate intervals; each apex finely lunate and shortly bispinose, latero-apical
margins very minutely serrulate. Underside with fine, sparse, recumbent pile;
prosternum strongly, metasternum moderately, abdomen minutely, punctate.
Dimensions: 9-10 « 3-2-3-5 mm.

Habitat: Victoria: Loch (A. H. Westley) in Coll. Wilson.

Two examples (? § ¢) sent by Mr. F. E. Wilson are, I believe, clearly distinct
from any described species. Of the form of S. titania, Cart., and S. nanula Kerr., by
its elytral pattern it would be placed near 9. horni Kerr. (unimaculata Cart.) in
my tabulation; though this is much larger and different in other characters. It is

BY H. J. CARTER. 227

narrower and more elongate than S. balteata Saund. The name fossoria (a digger)
is suggested by the spade-like marking of the elytra. Holotype (? ¢) in Coll.
Wilson.

STIGMODERA (CASTIARINA) DILATATA, nN. Sp. (Text-figure 5.)

Subcylindric; head and pronotum blue-black, scutellum violet, underside biue,
elytra red with the following markings blue; narrow basal border, wide premedial
fascia extended laterally forward but not reaching margins, an irregular post-
medial fascia, widened on suture, not connected with the former but connected
along suture with large, sagittate, apical mark, the point of the arrow blunted
and widened at extreme apex. Head deeply excavate and channelled between
eyes, strongly and closely punctate. Prothorax very convex, widest in front of
middle, apex advanced in middle and at sides, base bisinuate, sides widely rounded
and strongly dilated anteriorly; front angles obtuse, hind acute; disc with clear,
round punctures, fine and close anteriorly, larger and more distant behind,
slightly rugose at sides, medial line finely indicated throughout, medio-basal fovea
indistinct. Scutellum concave and laevigate. Hlytra as wide as prothorax at base,
sides nearly straight, apices with oblique excision not, or scarcely, bispinose,
subapical margins entire; striate-punctate, the seriate punctures small and almost
hidden in striae, intervals flat (except towards apex) and finely but clearly
punctate; underside closely punctate, with sparse, short pubescence. Dimensions:
12-125 x 4 mm.

Habitat.—Western Australia; Wembley Park (Mr. H. W. Brown).

Another of this keen collector’s discoveries, of which he has kindly sent three
examples. While allied to S. trifasciata Saund., and S. recta in form and elytral
pattern it differs from all of this group in the unusual form of the prothorax,
which is widely dilated in front of the middle, besides showing colour differences
from the above-mentioned.

Holotype (?2) in Coll. Carter.

STIGMODERA (CASTIARINA) FLAVO-VIRIDIS, n. sp. (Text-figure 6.)

Oblong ovate; head, pronotum, underside, antennae and femora metallic
green, elytra chiefly green, with the following markings yellow; a small latero-
humeral spot, a narrow, straight vitta outside the third interval, extending from
behind the base to the middle, here joining a narrow, straight fascia extending to
(and slightly widening at) the sides, and a short narrow, undulate fascia at the
apical fourth, this widely interrupted at the suture and sometimes (in four out of
eight examples) extending to the sides; tibiae and tarsi blue.

Head channelled and widely excavated, closely but not uniformly punctate.
Prothoraz rather strongly convex, widest at middle, apex sub-truncate, base
bisinuate, sides well rounded, arcuately narrowed in front, anterior angles obtuse,
scarcely sinuate behind, posterior angles acute; disc densely punctate, the punctures
becoming coarser, with rugose surface at sides; a finely cut medial sulcus
throughout, terminating in a basal fovea. Scutellum subcordate, concave and
punctate. Hlytra sinuate at sides, slightly widest behind middle, their apical
region finely serrulate, apex with a defined lunation, each apex finely bispinose,
sutural spine very short; striate, both intervals and striae alike closely covered
with punctures (of which no defined series is traceable), these becoming coarser,
with transverse rugosity, towards the sides; intervals convex, the third interval
strongly, the fifth less strongly so, these two wider and more raised than the rest.

228 AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES, V,

Prosternum rather coarsely, the rest of underside more finely punctate; abdomen
with thin clothing of recumbent, whitish hair. Dimensions: 10-12 x 4-4-5 mm.

Habitat—New South Wales: Mt. Kosciusko (Professor L. Harrison and the
author). Hight examples before me, seven taken by Professor Harrison in
February, 1927, on dwarf flowers (Cassinia, sp.), one taken by myself in
February, 1926. Wither all the examples are ?9 or there is no sexual dimorphism
of the abdominal segments. The only slight colour variation is a tendency for
green parts ‘to become blue in three examples (a common form of colour variation
in Buprestidae). In one example the elytra show purple patches. The species is
structurally allied to the S. serplagiata L. and G. and S. kershawi Cart., group,
in which one or two elytral intervals are more convex than the rest, though it is
more robust and parallel than these two. By colour it stands near S. wilsoni
Saund., and S. flavo-picta Boisd., the yellow markings being much narrower than
in typical examples of these species. Some Tasmanian examples of S. flavo-picta
Boisd., come rather close to it in colour, but are readily differentiated by the
more slender and sinuate form and the much finer elytral sculpture. S. dimidiata
Cart., is without any longitudinal vitta, besides having a less strongly sculptured
upper surface. Holotype, sex doubtful, in Coll. Carter.

NEOSPADES NIVEO-SPARSA, Tl. SD.

6. Oblong, attenuate behind, head coppery violet, pronotum dark bronze, elytra
bronze-brown irregularly marked with small spots of white pubescence, underside
metallic with a purple tinge, legs greenish, tarsi red, antennae coppery.

Head strongly, rather closely punctate, forehead with a medial carina, eyes
prominent (extending beyond the lateral border of pronotum). Prothorax: apex
subtruncate, base bisinuate, sides arcuately narrowed from base to apex, lateral
carinae diverging towards the front, the upper continuous throughout; two large
foveae on each side of middle anda pubescent depression above, and following the
line of, the upper lateral carina; disc transversely striolate, rugose punctate at
sides. Seutellum transversely ovate, with some 8-10 large punctures. EHlytra sub-
parallel for two-thirds of length, thence obliquely narrowed, apices separately
rounded, and fringed with bristly spinules; scalose punctate, the pubescent spots
more evident along suture and sides; segments of abdomen with pubescent patches
at sides; sternal regions and legs also finely pubescent; external margins of hind
tibiae with three elevations each bearing a pencil of spinules near the apex.
Dimensions: 11 x 4 (+) mm.

Habitat—New South Wales: Bogan River (Mr. J. Armstrong).

I am indebted to the above young naturalist for the capture of this distinct
species. It should stand near N. gouldi Hope in my table (These Procrrpines, 1923,
p. 172), from which it is abundantly distinguished by size, form and colour. The
outline is somewhat like that of Cisseis careniceps as depicted in Fig. 8, p. 169 of
my Revision (These Proceepines, 1923). The tibial elevations are very much
as in WN. lateralis Blkb., except that the third (nearest apex) is continuous between
the second and the apex, whereas the first and second are isolated humps.
Holotype (sex doubtful) in Coll. Carter.

NEOSPADES CUPRICAUDA, 0. SD.
g. Elongate oblong, head and pronotum golden green or fiery coppery; elytra
more or less violaceous or cyaneous, with coppery chiefly at base, sides and apex—
the limits of these colours variable and vague—with four pubescent spots towards

BY H. J. CARTER. 229

apex in an arcuate row, one on each side about half-way, and a few others
irregularly dispersed; sternal regions and legs generally greenish, abdomen fiery
copper.

Head rather strongly grooved between the eyes, regularly and strongly
punctate (less densely than in N. lateralis Blkb.). Prothorax: apex and base
bisinuate, sides nearly parallel on basal half, arcuately narrowed in front, posterior
angles subrectangular, the lateral carinae meeting near these angles, parallel for
the greater part, slightly diverging at front; disc with moderately light, shallow
punctures, with only a slight tendency to transverse strigosity at sides, depressed
on each side towards base. Scutellum very transverse, with a few large punctures.
Elytra of same width as prothorax at base, lightly compressed behind shoulders,
subapical margins finely serrulate; surface finely scalose punctate; prosternum
densely and deeply punctured, metasternum and basal segment of abdomen
moderately punctate, the remaining segments finely and much more sparsely so.

°. Differs in larger size and more obscure colour; the head, pronotum and
the whole underside (including legs) being dark copper bronze, elytra largely
cyaneous bronze at base and sides, generally coppery at extreme apex. Dimensions:
7-9 x 2-5-3 mm.

Habitat—Northern Territory: King River (Sir Baldwin Spencer).

Mr. J. A. Kershaw has sent twenty-eight (19 64, 9 22) of this species from
the National Museum for determination. It is in size and colour intermediate
between N. chrysopygia Germ., and N. lateralis Blkb., the colour definition being
less regular than in either; but it differs strongly from both in the finer sculpture
of both pronotum and elytra—the former having little transverse rugosity, the
latter a much less deeply impressed surface. The pronotal structure is also
intermediate—less convex than with lateralis; more convex with sides less
explanate than with chrysopygia. The pronounced sexual colouration may be
noted as a further distinction.

Holotype ¢ and allotype 9 in National Museum, Melbourne.

CISSEIS SUB-BIFASCIALIS, N. Sp.

Oblong, little attenuate behind; head coppery, pronotum greenish bronze with
white pubescence; elytra blue with a preapical fascia and a half-formed post-
medial fascia of white pubescence; underside coppery bronze, abdomen with
lateral patches of pubescence, appendages coppery.

Head densely punctate, with well defined medial depression, eyes not
prominent, head (including eyes) not wider than apex of prothorax. Prothoraz:
apex and base bisinuate, sides nearly straight, feebly widened near base, here
both lateral carinae visible from above, these sub-parallel, rather close and
continuous to apex; disc largely covered with silvery pubescence, leaving the
medial and apical area bare, with fine striolate-punctate surface. Elytra oblong,
lightly enlarged at shoulders, and compressed at middle, apices widely and
separately rounded and minutely spinuled at margins; disc scalose-punctate,
sparsely pubescent near base besides on the fasciae, the post-medial fascia
represented only near sides, extending thence about one-third of the way across,
the preapical fascia uninterrupted; underside finely and closely punctate.
Dimensions: 6 x 2 mm.

Habitat—New South Wales: Bogan River (Mr. J. Armstrong).

Another of Mr. Armstrong’s discoveries shows a species that resembles in
colour and clothing the well-known Hypocisseis suturalis Saund., but differs from

230 AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES, V,

that species in its narrow, sub-parallel form and the structure of the head and
prothorax, the head being of the normal Cisseis type. It belongs to my Section II,
Group B. Holotype (sex ?) in Coll. Carter.

TENEBRIONIDAE.

Pezohelaeus denticollis Cart. Mr. Hacker has lately taken this on Tambourine
Mountain, Queensland, and sent it to me for determination. Like the unique type
(in the Melbourne Museum) it is a ¢. I now think it possible that this is the
male form of P. hirtus Macl., of which I only know the female. The two appear
identical in form and sculpture, differing markedly in the anterior angles of the
prothorax—strongly dentate in denticollis—rounded with a slight posterior
Sinuation in hirtus. Pezohelaeus should, I think, rank as a subgenus of
Pterohelaeus, distinguished only by its enlarged tarsi. The sexual character
suggested above has its analogy in Anausis metallescens Westw. (see These
PROCEEDINGS, 1913, p. 71) though absent from the eighty-two known species of
Pterohelaeus. P. hirtus Macl., is rare in eastern New South Wales—Strathfield
(H. Cox), Kandos (T. G. Sloane), Gingken (R. B. Carter) being localities recorded
by me.

Pterohelaeus guerini Br. = P. bullatus Pasc. = P. ventralis Blkb. = ? P. regularis
Blkb. = P. puer BIkb.

Having. collected long series from New South Wales, Queensland, N.W.
Victoria and various parts of Western Australia, I am now convinced of the
above synonymy. Champion recorded P. bullatus Pasc., from Fremantle, W.A.,
when describing the Australian collections of Commander J. J. Walker, M.A.,
R.N. (Trans. Ent. Soc. London, 1894, p. 383). I have seen the type of P. regularis
in the British Museum and a cotype in the South Australian Museum of which
my notes say, “very doubtfully distinct from bullatus—a common species in the
Narrabri district of New South Wales’. A cotype of P. puer has recently been
sent for examination from the South Australian Museum.

LICINOMA.

Since publishing my tabulation of this genus (These ProcrEpines, 1920, p. 243)
I have added eight new names. I therefore append a revised tabulation.

Licinoma.
ieeMiytransenriate- OLestriate-punGtate 1. cm ccine eis anions Se ich hare Pea eee Penarth ee 2
Hilytra striate or SUICate monly? Feateecccc: ever Mtoe re Voteceue ee oee ne anes corrals se en HHO M ERR Teil e eva eee meee 22
2. Elytral intervals of uniform width .............; MIS TOIL BIS nto cio tb. Sees hone o 3
Miyvinal intervals Not OLeinitornay wa dith yy cuciec is sqsnsisteecoueletiel oheeielelcmenekemene: oilicnse sera sicus 19
3) SESLYVETaAL | INTERVALS! Tate Ae raretoe ap ose seh o Sere rao abe ve HMA ie cide IRCA aL eas apt OREM RRR acre alyssa eh a 4
Bivtral: intervals (CONVEX ice ei cuseice Geissler dabetecetaascebons obelie) eben al Monee Mesmensuaeel hel /oereu oo) all
4 ,Elytrav intervals» clearly Dunetate sie cals cue e cere ee eT etenedeliehememaisieteltorehens veiteptaus tarnene 5
Hlytiralwntervals: ini prim Crate: we cassette tece meteor oheie bee ialieta etehotche ke Men eN feu eneiestel oleh dey el Susiekens 8
Ie) SELOLHOLAX I tUANSVeELSe, ISIGESH Well TOUNGEN pane iacbaie lionel ieeekedenensiencao tis ielapelieteleeie 6
Prothorax as lone asi wide, sides mearly, Stralentera scien ae eie ie iaiciche cileienetecencns 1
Gs PErouOLuImsmOreuorn Ess channelled ia. caer ertie mies niet ele nela subcanaliculata Cart.
EVOUOtUM HOt TCU ANTICS foie ee atosnete cieteu diets le tacn, heals tate eieeKnenete tober sits rials nitida Pase.
isiaslytral intervals imoduloseiy:).;syastetethe dete cies cispake obdte ts tepapahenomeltaheretante ys nodulosa Champ.
Hilytral intervals not nodulose’ 252 fare sees es Jails ater aUstelts Wilotests Rate Pekeel 3 tasmanica Champ.
S$. “Hiyiral untervals Modwlosey Ailes etleveteusie thas ous oie lo aupeshonsmetarencuanmlonisiens umbilicata Cart.
Dlytral AntenVvals NOte TOUAULOSE: cs aaisui incest) stein HOt IC neiei lic icivcseley ckeacus Tsu edeucteieuci 9
DO, TAS TAC ite NS he ee oi oe ET Cae TEE Bale oes tol SV COMM on ees UcUsbomey mic eicension eccits, siiatehetniovene 10

DESEEESTACLOUS IA sie ere « bp Bia ode ae ee le tone aoe laa DUaTe PEN MORO e Ret ohe lotebaliol ara toN Nees pallipes Blkb.

BY H. J. CARTER. ' 231

{0eeLbindsanslestol pronotumy rectan ewan cusses oiler enalenetelicueiallasiereher siete nitidissima Lea
LMndeanSesTorspronoOLctum TOD EUSE ms sccreienetl ene siete clclstelysusieysucie) seleed el len eneie planata Cart.
Mbt ee Ey ET ae LUC unetcien eur teres cas cn. (ah vicncl cnemed SreM enn eae neo eoneraue lala) olen abicnale id cusienepelamtanstareyersuateres ore 12
Mi lyitrarnDlackVor i DEOMZEM te. cscs ehecehedeasettmenerenctenerie(to) oe res vette: Scie) eiay seit epee auction ate omelette 13
12 uPronotum Ccoppery, without medial ichammnel Wen: mis - 2 lel ere se hela aie splendens Cart.
IPO OW ban, LMS: \yiAtela weVEChiEy| oe ooaodosnoadooonsaanensooucsnonUe coerulea Cart.
PS EP rOCh OLAxe ELAN SVCESC fasts; sien suehish sicusuelewe dememeueneu shave tele usie csi elscist ate sitay cu sue veitelsatanelalior sueulap ence cnet 14
IFTLOLIOna On Ser chaniawilde cimiieierreacicierettsrsiere tans nou cue cuca ele oueiaiecerciiel meridiana Cart.
aE LOLIOLAS LWLGCES ty at ILO GLE Ai ak roeeeoenetene etestelen cilokene foie lerietiarieccl seal ater aialouttete ran nae cae 15
Prothora xawilGest pny front 0 f end Ce miyeperciiny kari stsicne chor clel seshaievchocieictcnemebe enone alana 107/
IL; JAeCmMOikinn Shicxokcobllere havel BONN AWHEENKS Gobcuaucosdudgoaoanobon4oe cyclocollis. Cart.
IPROMNO MHI. MONE Kibo nMlene Olicehalhy jRUNNCURNH Sobancubuonbnosoonobcduocnosuouuue 16
1G, Iba MASAI meENATON CHOC IOWIKCUEI®. cotboucccdnonaoo ds odcdbnnoaumes elata Pasc.
milytral intervals widevand! striolavelomlyaycir «ey. e) > vlebe « seiele o elNel shee gilesi Cart.
U7/. TiopOrwbon Wray Taschen Oech Godiooddgocbunudaccongoandcouuunact's 5 truncata Cart.
Pronotume withoutenne dial 5 chiaimi ell aero sesess eyes cieien ene sicyets, cher oj susie) meme ene ey alicia oe 18
See teadeandmpronotum: Strong lya pum Ctalemmcricretepeienct ss) cucdehoiaie-baieleieie monticola Blkb.
Eeadvand pronotum very, finely wunectate = a... scm sce sie sles sere apasioides Cart.
Oy bral mbervials: flat hi Li. vical nadeweme dene mat names ra vebicb stiaetrerlcnrewoeeilcy ehialvoucnonsteclioperonc vote ube ae 20
Hyer wa ntervall SivGOmV exe oii: evyy A lcpentane Menem than wanet od ots dalled op sls ocgs. oi eeevs hen Ausirebcttay cas aerea Cart.
YO weeiecadvuandspronotumicoanrseliy: puUNnChaley ciuyer-wienel seis cieia)e! ceed ch ele: aust puncticeps Cart.
Eicadwandapronotiimy» vVieny - nel ys pln CuaLe maesrse-i eit aie slic) eiersie) <usi stele sie) saeucten serene ais PALL
Qieeivoraleincenvals 1mMpunctate ang snNOM-=SCLOSS a) .tes eee er. cilia ajietelalcnelonel cite sylvicola Blkb.
Elytral intervals punctate (third and fifth setose) ................ commoda Pasc.
PO. Jdibyoreyl soy ChE babbopMaDy AKO “so do cangsoacasonougcdobasogauGonbouooEdor 23
DIA TEMECIAVEUIS) sae CRE Whoubreyaon WVU sosoooogones FbogsuudodoooUdEOS illidgei Cart.
23. Colour violet-bronze, pronotum with medial channel ............ angusticollis Cart.
Colour black, pronotum without medial channel ...................... major Cart.

ENHYPNON PUNCTATUM, D. SD.

Obovate, very convex, opaque brown, antennae, tibiae and tarsi red, clothed
extensively with rough hairy derm, becoming bristles on the legs.

Head less wide in proportion to pronotum than in #H. laticeps but similar in
form; eyes prominent, antennae, first segment very stout, 2 elongate subconic,
8 shorter and narrower than 2, 3-9 subconic, gradually increasing in width, 10-11
clavate, 11th elongate ovate, twice as long as 10. Prothoraxr very convex, base
truncate, apex with bilobed process overhanging the head, widely excised between
these lobes, sides involute, margins unseen from above, disc with two well raised
ridges (forming an extension of the apical lobes), occupying a great part of
surface; basal third and sides covered with small tubercles, the ridges with felt-
like clothing. Hlytra of same width as prothorax at base, humeri obtuse, the
outline subhexagonally obovate, widening at first obliquely, then angulately
subhorizontal, then angulately narrowing towards apex; surface nodulose with
two elongate ridges, one on each side of scutellum, extending from the base a
short distance, two blunt tubercles behind each of these en echelon, four smaller
tubercles in a transverse line near middle, a third group of about six tubercles on
apical declivity and smaller ones behind these near apex. The area unoccupied
by tubercles with regular series of large round punctures, the two sutural rows
uninterrupted. Dimensions: 5 (vix) x 2 (+4) mm.

Habitat—Tasmania: Mount Wellington (H. H. D. Griffith).

A specimen, labelled as above, has been kindly sent me by Mr. A. H. Elston.
It is much larger, with more pronounced excrescences, than EH. laticeps Cart.,
the only other member of the genus, and is easily distinguished by the large seriate
punctures of the elytra, which are either nonexistent or covered by derm in
laticeps. It is doubtless one of the moss-dwelling Coleoptera that are so numerous
and interesting in the mountain fauna of Tasmania. Holotype in Coll. Carter.

232 AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES, V,

NYCTOZOILUS TAYLORI, nN. Sp.

Oval, convex, subnitid black, glabrous.

Head densely, finely punctate; antennae with 38 longer than 4-5 combined,
8-10 round, successively diminishing in size, 11 pyriform. Prothoraz, 5 x 8-5, very
convex, apex arcuate-emarginate, base nearly straight except for produced angles.
Sides widely and evenly rounded, anterior angles subacute (slightly blunted at
tips), border with widely rounded reflexed rib, a deep and rather wide convexity
between border and disc, hind angles clearly dentate, pointing backwards; disc
with indications of medial channel, a foveate depression at middle near base,
whole surface with close, fine, shallow punctures. Scutellum very transverse.
Elytra widely ovate, each with three nitid, undulate costae, the first two connected
near apex, the intervals irregularly subfoveate with ill-defined reticulations,
becoming obsolete at sides and apex. Prosternum minutely striolate, abdominal
apical segments minutely punctate. Dimensions: 19 x 11 mm.

Habitat.— Queensland: Townsville (Mr. F. H. Taylor).

A single 9 example kindly sent by Mr. Taylor, entomologist of the Institute
of Tropical Medicine. It is nearest to N. carlovillensis Cart., but differs in larger
size, more nitid surface, more convex pronotum, with wider ribbed border, elytral
reticulation less elevated and defined. Holotype in Coll. Carter.

CAMPOLENE CYANEA, 0. SD.

Oblong, oval, convex, peacock-blue, nitid, antennae and legs red, underside
nitid, black, glabrous.

Head and pronotum coarsely and rather closely punctate, with a subfoveate
impression in front of eyes. Prothorax convex, sub-gibbous at middle near apex,
widest near front, base feebly bisinuate, sides nearly straight, with a narrow
horizontal border. EHlytra of same width as prothorax at base, striate-punctate,
the punctures in striae larger than in C. nitida Pasc., intervals convex and
laevigate. Underside even more differentiated from C. nitida by the coarse
punctures of the prosternum and abdomen, those on the latter somewhat irregular
in size and position. Dimensions: 7-9 x 3-3-5 mm.

Habitat.—Queensland: Dalveen (the author and R. Illidge); Stanthorpe (Mr.
F. A. Perkins and L. F. Von Wieldt).

Seven specimens examined show a structure very similar to that of the well-
known C. nitida Pasc., but differing in colour, the more gibbous pronotum, and
the much stronger sculpture of the upper and lower surface. I have long had a
Single specimen under observation and the acquisition of further material from
Stanthorpe shows constant distinction from Pascoe’s species. Holotype in Coll.
Carter.

Cardiothoraz consimilis Cart. (1926) = C. australis Cart. (1906) var. With
further material I do not consider that the distinction of this species from
C. australis can be maintained. Mr. Hopson informs me that “southern plateau”
meant southern highlands of New South Wales—Moruya region—so that my
locality (Barrington) should be corrected.

CISTELIDAE.

I took a number of Hemicistela testacea Cart., by beating the flowers of
Banksia sp., at Geraldton, Kojarena and Eradu (W.A.) in September, 1926, in
company with Amphichroum sp. In this genus there appear to be two lamellae
on each of the anterior and only one on each of the four posterior tarsi (or if

BY H. J. CARTER. 233

two be present on the intermediate tarsi one is so small as not to be evident with
a Zeiss binocular); whereas in the majority of Australian Cistelidae the formula
for such lamellae would be 2, 2, 1 as stated by me (Proc. Roy. Soc. Vict., 1915,
p. 52).

Allecula morio F.—Specimens of this from South France have been sent
me by Mr. K. G. Blair. I find two lamellae on each of the four anterior tarsi
and one on each posterior tarsus. This is inconsistent with the classification of
Solier and Mulsant for this genus ‘‘one small lamella on the penultimate joint of
each tarsus’”’.

ANAXO CYLINDRICOLLIS, Nn. Sp.

9. Elongate, oblong, nitid, bronzy black, rather thickly pilose as to upper
surface and appendages, more sparsely so below, with suberect white hair; tibiae
and basal two-thirds of femora red, its apical third black; antennae opaque,
appearing greyish from their white pubescent clothing; tarsi dark.

Head densely punctate, strongly pilose, eyes widely separated, antennal
segment 3 longer than 4, 4-10 subequal, each slightly enlarged at apex, 11th very
short and oval. Prothoraz longer than wide, nearly cylindrical, widest near front,
lightly narrowed towards base; apex subtruncate, base feebly produced in middle,
lateral border and angles unseen from above; disc strongly, not very closely,
punctate, a narrow medial channel on basal third and slightly flattened in this
region; otherwise with strong transverse convexity; pilose at sides. Scutellum
semicircular. Hlytra nearly twice as wide, and two and a half times as long as
prothorax, sides parallel, shoulders rather square; striate-punctate, the striae
shallow, seriate punctures larger than in A. cylindricus Germ., intervals flat
and everywhere transversely rugose; clothing subrecumbent near suture, upright
at sides. Underside finely and sparsely punctate and pilose. Dimensions:
10 x 3 (+) mm.

Habitat—Western Australia: Bridgetown (the author).

6. Wanting. I took the unique example in August, 1926, when visiting
the south-west with the members of the Australasian Association for the Advance-
ment of Science Congress. It is clearly distinct from its nearest ally, A. dentipes
Cart., in which the elytral intervals are almost smooth and the sternum densely
pilose (almost glabrous in cylindricollis). A. cylindricus Germ., is nitid black,
with a wider, less convex, and more. finely sculptured prothorax, and convex
elytral interstices. Holotype 9 in Coll. Carter.

HYBRENIA ILLIDGEI, n. Sp.

Elongate oblong, nitid black, body and legs sparsely (rather densely on
pronotum) clothed with long, upright, black hair.

Head rather coarsely and sparsely punctate, eyes large, prominent, separated
by a space of about half the diameter of one eye, antennae with third segment
linear, 4-10 subtriangular and subequal, 11th ovate-acuminate. Prothorax rather
convex, apex slightly advanced in middle, anterior angles depressed and blunt,
base gently bisinuate, posterior angles sharply rectangular, sides parallel on basal
half, rounded in front, disc with sparse and irregular coarse punctures, each
bearing an upright hair, a wide medial depression and a round foveate depression
near hind angles. Scutellum large, almost smooth, rounded behind. JElytra
considerably wider than prothorax at base, shoulders prominent but rounded,
sides parallel; striate-punctate, the punctures in striae large, square and close;

234 AUSTRALIAN COLEOPTERA: NOTES AND NEW SPECIES, V,

intervals rather sharply convex, the suture itself subcarinate behind a well-marked .
post-scutellary depression; each interval with a single line of small setiferous
punctures; sterna and epipleurae coarsely punctate, abdomen much more finely so;
post tarsi having first segment as long as the rest combined. Dimensions:
12-13 x 4-4-5 mm.

Habitat.—Queensland: Brisbane (the author), Queensland National Park
(Mr. A. Musgrave). d latet.

Two female examples examined show a rather narrow, parallel species with
the general facies of H. angustata Macl., but easily distinguished from this and
other species having convex elytral intervals by the combination of nitid and
strongly pilose surface, sparsely and coarsely punctate pronotum, the elytral
seriate punctures of the same average size as in H. angustata Macl.* From
H. pilosa Cart., it is distinguished by its parallel form, black hairs, much more
sparsely punctate pronotum and the elytral intervals without wrinkles. Holotype
in Coll. Carter.

CHRAMBYCIDAH.

Microtragus—When I wrote my Revision of Athemistus and Microtragus
(These Proceepines, 1926, p. 492) I was unaware of the paper by Professor
Aurivillius (Arkiv. for Zoologi, 1917) on the Cerambycidae of the Mjoberg
Expedition, a copy of which has lately come into my hands. In this paper three
species of the above groups are described as new—(a) Microtragus cristulatus,
(6) Athemistus (Hoplathemistus) conifer and (c) Athemistus (Hoplathemistus)
albofasciatus—and a new subgenus created. I think M. cristulatus is one of the
many varieties (var. sticticus Pase.) of M. arachne Pase. almost black, that were
amongst those that I examined from the National Museum. The proposed new
subgenus Hoplathemistus would appear to include my group B of Microtragus, if
the antennal sockets are approximate. Unfortunately Professor Aurivillius makes
no mention of this important character, which together with the presence of the
humeral spines I take to be the main distinction of Microtragus from Athemistus.
If this conjecture be correct my M. echinatus may be synonymous with
A. (Hoplathemistus) conifer Auriv. The affinities of Hoplathemistus are, I think,
with Microtragus, of which it is a subgenus, rather than with Athemistus.

*In H. angustata Macl., the medial striae are almost impunctate, while the lateral
striae contain very large punctures: the surface is glabrous.

‘

THE GASTEROMYCETES OF AUSTRALASIA.
vii. Tue GENERA DISCISEDA AND ABSTOMA.

By G. H. CunNINGHAM, Government Mycologist, Department of Agriculture,
Wellington, N.Z.

(Plates xx-xxii.)
[Read 29th June, 1927.]

In 1892 Morgan separated from Bovista species possessing a capillitium of
free, short, simple fragments of hyphae and placed them in a genus he named
Catastoma. In the type species ef this genus, he noted that the stoma was at
the base of the plant, and made this one of the chief characters of the genus.
Since the publication of this paper many species have been recognized, and it is
now known that the position of the stoma is of no generic value, for it may be
apical or basal.

The genus was separated many years previously by Czernaiaiev (1845), who
named it Disciseda, but his paper was overlooked until Hollos (1902), from a
perusal of Morgan’s paper was able to recognize the fact that Catastoma was the
same as Disciseda. Lloyd in his publications has taken exception to the use
of Disciseda, and throughout uses Catastoma, a name which is untenable according
to the Rules of Nomenclature.

The species are all similar externally, and in consequence difficult of
delimitation. This difficulty has been increased because many species have been
named and imperfectly described from single specimens.

Lloyd (Myc. Notes, 1922, p. 1167) groups the species according to the colour
of the gleba, but this is impracticable since in many the colour is not constant,
different plants in the same collection, agreeing in all other respects, possessing
an olivaceous, umber or purplish gleba (D. candida, D. cervina and D. hyalothriz).

After extended examination of the collections enumerated in this paper, the
writer is of the opinion that two groups of characters alone, those of stoma and
spores, may be used for the separation of the species.

The stoma may be a simple, plane, poorly defined aperture (D. pedicellata,
D. hyalothrix); or may be conspicuous, mammose and fimbriate (D. cervina,
D. hypogaea); lacerate and toothed (D. verrucosa); or definitely tubular and
surrounded by a depressed groove (D. anomala and D. australis). As in old and
weathered specimens these characters may be obliterated, the spore characters,
which do not change with age, will be found to be more definite and accurate.
The genus may in this manner be divided into pedicellate and apedicellate sections;
and further separation may be effected according to the nature and to the degree
of roughness of the spore-markings (see key).

Abstoma is separated because of the absence of a definite stoma, dehiscence
being effected by the irregular rupture of the peridium. One other character of

D

236 GASTEROMYCETES OF AUSTRALASIA, Vii,

interest in this genus is the fact that in the two species known, the spores are
strongly reticulate.

Members of both genera consist of a peridium enclosing the gleba. The
peridium consists of the usual two layers, exoperidium and endoperidium. The
exoperidium is in most species in the nature of a membrane composed of hyphae
and sand grains or vegetable debris immixed, forming a firm but brittle structure
which may attain a thickness of 2-3 mm. or more. As the plant matures this
usually fragments and falls away, save for a persistent discoid or cupulate basal
portion. In some species the exoperidium is membranous.

The endoperidium is membranous or papyraceous, tough and usually polished,
though in one or two species it is furfuraceous. It is perforated with a solitary
apical or basal stoma. In one or two species a third delicate gelatinous and
reticulate membrane is sometimes present lying between the layers of the
peridium. As it may be present or absent on different plants of the same collection,
it cannot be considered as being of specific value.

The gleba consists of capillitium and spores. It is usually pulverulent, but
is compact in Abstoma purpureum. The threads are usually short, 50-100 » or
more, usually coloured, simple or short-branched, straight or more frequently
curved, continuous and non-pitted. The spores are continuous, coloured, pedicellate
or apedicellate, variously roughened, and usually globose. No sterile base or
rooting base is present.

The habit is noteworthy, most species being hypogaean, growing solitary
or in small groups in the ground, and becoming exposed as a rule only at
maturity.

All descriptions are original and have been drawn up from specimens examined
by the writer.

The writer is again indebted to Dr. J. B. Cleland, University, Adelaide, for
the loan of all specimens in his possession. :

DISCISEDA Czernaiaiev.

Bull. Soc. Imp. de Moscou, 1845, xviii, p. 153—Catastoma Morgan, Jour. Cin.
Soc. Nat. Hist., 1892, xiv, p. 142.

Peridium depressed-globose, of two layers; an exoperidium which may be
thick or thin, membranous, or compact when formed of hyphae immixed with
sand particles or vegetable debris, fragile, breaking away irregularly, save a
small discoid or cupulate basal portion; endoperidium papyraceous or membranous,
tough, variously coloured, smooth or furfuraceous, dehiscing by a definite stoma,
which may be apical or basal; sterile base absent.

Gleba pulverulent; capillitium of short, simple or short-branched, continuous,
non-pitted coloured hyphae. Spores globose, coloured, variously roughened,
pedicellate, apiculate or apedicellate.

Habitat:—Solitary or in small groups on or in the ground.

Distribution:—Europe; Asia; Africa; North and South America; Australia;
New Zealand.

A small genus of about twelve species, eight of which occur in Australia and
New Zealand.

Key to the species.
Spores long-pedicellate (10 w or more).
Spores strongly verrucose, 8-10 w diam. ..............2+++.+> 1. D. pedicellata.
Spores finely verrucose-areolate, 10-13 wi... . cece e eee e ee enee 2. D. hyalothria.

BY G. H. CUNNINGHAM. 237

Spores apedicellate (or apiculate, stumps only of the pedicels

persisting).
SDOLESPalMOSEISMOOCMW raise ccs scorsrsueuel ena kone uel sists ails) eves aia: eu susuetione sive 3. D. candida.
Spores finely verrucose.
Stomay imibriate=m'amim OSewy einer neleciclcienteroies scaielle cielans 4. D.cervina.
Stoma tubular, surrounded by a depressed groove.
Spores finely verrucose-echinulate .................. 5. D. anomala.
Spores covered with fine round-topped warts ....... 6. D. australis.
Spores coarsely verrucose.
Spores covered with flat-topped echinulae ............... 7.. D. hypogaea.
Spores covered with coarse verrucae ..............--.-- 8. D. verrucosa.

1. DIScISEDA PEDICELLATA (Morgan) Hollos. Plate xx, figs. 1, 2; Plate xxi, fig. 4.

Term. Feuz., xxv, 1902, p. 103—Catastoma pedicellata Morg., Jour. Cin. Soc.
Nat. Hist., 1892, xiv, p. 148.

Peridium up to 3 cm. diam., depressed-globose or lenticular, attached by a
small rgoting base; exoperidium a thick sand-case, of hyphae and debris immixed,
grey or brown, flaking away save a small discoid basal portion; endoperidium
tough, membranous, chestnut brown or umber, smooth, shining, dehiscing by a
small apical, plane stoma.

Gleba purplish, pulverulent; capillitium pallid chestnut, of the usual type.
Spores globose, 8-10 uw diam. (including verrucae), pedicellate, pedicels up to 25 u
long, stout, tinted; epispore chestnut brown, coarsely and strongly verrucose or
verrucose-echinulate.

Habitat:—Solitary on the ground.

Distribution:—North America; South Africa; Australia.

New South Wales:—*Sussex Island, Clarence River, Sept., 1909; “Narrabri,
June, 1919 (Det. Lloyd No. 527 as Catastoma pedicellata).

South Australia:—*Yanco Area, Nov., 1919; *Pt. Hlliot, Aug., 1918, D. I.
Cleland; *Buckland Park, Apl., 1923; *Fullarton, Adelaide, May, 1923; *no locality,
F. R. Zeitz.

The species is characterized by the (usually) large size, firm, leathery, umber,
polished endoperidium, indefinite plane stoma, and especially by the long-
pedicellate, coarsely roughened spores. The spore markings vary somewhat. in
different collections; in some being strongly verrucose, in others verrucose-
echinulate. The length of the spines may vary from 1:5 uw to 0:5 uw. As all
conditions may be noted in the same microscopic field, these variations cannot be
considered of specific import.

This is one of the most abundant of the Australian species, judging from
the collections in Dr. Cleland’s possession, and is separated from D. hyalothriz,
which it resembles in the pedicellate spores and plane stoma, by the smaller,
more coarsely warted spores and longer pedicels.

2. DISCISEDA HYALOTHRIX (Cooke and Massee) Hollos. Plate xxi, fig. 5. ~

Noev. Koezl., 1902, p. 107.—Bovista hyalothriz Cke. et Mass., Grev., 1888, xvi,
p. 73.—Catastoma hyalothrix (Cke. et Mass.) Lloyd, Lyc. Aus., 1905, p. 27.

Peridium up to 2:5 cm. diam., depressed-globose; exoperidium in the nature
of a sand-case, flaking away irregularly, save a small attached basal portion;
endoperidium umber or purplish, smooth, tough, membranous, dehiscing by an
irregular, plane, apical stoma.

* An asterisk denotes that the collection in question is in the herbarium of Dr. J. B.
Cleland, Adelaide; and where no collector is given, signifies that the collection was made
by Dr. Cleland himself.

238 GASTEROMYCETES OF AUSTRALASIA, Vii,

Gleba dark olivaceous, becoming dark purple, pulverulent; capillitium threads
tinted, of the usual type. Spores globose, 10-13 » diam., pedicels short, up to 15 yu,
stout, tinted; epispore deep chestnut brown, densely and finely covered with
tinted or hyaline verrucae, which appear areolate in consequence of their close
arrangement.

Habitat:—Solitary on the ground.

Distribution:—Australia.

Victoria:—Lake Allacutya, C. French. Type, in Herb. Kew.

New South Wales:—*Dubbo, June, July, Oct., 1915 (One collection det. by
Lloyd No. 137 as Catastoma hyalothriz).

South Australia: —*Adelaide, July, 1914.

Characterized by the pedicellate, closely and densely verrucose spores. The
verrucae are densely packed, and somewhat irregular in shape and size,
consequently in surface view the epispore appears somewhat areolate. The
plane stoma is also characteristic. The gleba may be purplish or olivactous.

3. DISCISEDA CANDIDA (Schweinitz) Lloyd. Plate xx, figs. 3, 4; Plate xxi, fig. 7.

Mycological Notes, 1902, p. 100.—Bovista candida Schw., Syn. Fung. Carol.,
No. 333, 1822.—B. circumscissa Berk. et Curt., Grev., 1873, p. 50.—Catastoma
circumscissa (Berk. et Curt.) Morg., Jour. Cin. Soc. Nat. Hist., 1892, xiv, p. 143.—
Disciseda circumscissa (Berk. et Curt.) Hollos, Term. Fewz., 1902, xxv, p. 102.

Peridium up to 3 cm. diam., depressed-globose; exoperidium thick, firm, of
hyphae and vegetable debris immixed, breaking away circumscissilely from the
base; endoperidium ferruginous to umber, tough, covered in part by a reticulate,
gelatinous layer; dehiscing by a basal, fimbriate, mammose stoma.

Gleba olivaceous, umber or purplish, pulverulent; capillitium threads pallid
chestnut, of the usual type. Spores globose, 3°8-4:5 u diam., with stumps only
of the pedicels remaining; epispore pallid chestnut brown, very finely verruculose,
almost smooth when ~x 1,000. :

Habitat:—Solitary or in small groups in pastures; hypogaean.

Distribution:—Hurope; North and South America; Australia; New Zealand.

New South Wales:—*Milson Island, Hawkesbury River, Feb., 1916 (two
collections, both det. by Lloyd No. 139 as Catastoma hyalothriz).

New Zealand:—Roxburgh, Otago, Aug., 1925, J. C. Neill.

Characterized by the almost smooth spores, absence of a definite pedicel, and
fimbriate, mammose stoma. According to Morgan the stoma is situated at the
base of the plant.

The gleba is olivaceous when young, deep umber when old; frequently in
Australian plants identical in all other respects, it is tinged with purple.

4. DISCISEDA CERVINA (Berkeley), n. comb. Plate xx, fig. 5; Plate xxii, fig. 1.

Bovista cervina Berk., Ann. Mag. Nat. Hist., 1842, ix, p. 447.—Globaria
debreciensis Hazsl., Zoo. Bot. Ges., 1877, xxvi, p. 226.—Bovista subterranea Peck,
Bot. Gaz., 1879, iv, p. 216.—B. argillacea Pat., Bull. Soc. Myc. Fr., 1888, p. 93.—
B. debreciensis (Hazsl.) de Toni, in Sace. Syll. Fung., 1888, vii, p. 476.—Catastoma
subterranea (Peck) Morg., Jour. Oin. Soc. Nat. Hist., 1892, xiv, p. 143.—
C. debreciense (Hazsl.) Hollos, Term. Koezl., 1902, lvi, p. 186.—Disciseda
debreciensis (Hazsl.) Hollos, Term. Feuz., 1902, xxv, p. 102.

BY G. H. CUNNINGHAM. 239

Peridium up to 4 cm. diam., depressed-globose; exoperidium thick, brittle,
flaking away irregularly save at the base; endoperidium tough, membranous,
purplish or tan coloured, furfuraceous, dehiscing by a fimbriate, mammose stoma.

Gleba olivaceous, umber oi purplish; capillitium threads pallid chestnut
brown, of the usual type. Spores globose, 5°5-6-5 » diam., with stumps only of
the pedicels remaining; epispore chestnut brown, closely and finely verruculose.

Habitat:—Solitary in the ground; hypogaean.

Distribution:—Europe; North America; Australia; New Zealand.

New South Wales:—*Forbes, Aug., 1915; *Merebine, Nov., 1918 (Det. Lloyd
No. 508 as “near Catastoma circumscissa’); *near Barellan, Aug., 1918.

South Australia:—*Grange, July, 1914; *Monarto South, July, 1922 (Det. Lloyd
No. 852 as Catastoma subterranea); *Ooldea, Aug., 1922.

New Zealand:—Roxburgh, Otago, Aug., 1925, J. C. Neill; Ashburton,
Canterbury, Aug., 1925, J. C. Neill.

The species resembles the preceding in many characters, but may be separated
by the larger and more definitely verruculose spores: The verrucae differ
considerably in different collections, some being fine, approaching those of the
former species (when the spore size alone separates them); others being coarse,
approaching those of D. anomala. In such cases the nature of the stoma aids in
separating the two.

Lloyd has examined the type of “Bovista cervina’ and found it to be the
same as “Catastoma subterranea’. Therefore the specific name cervina has
priority.

5. DISCISEDA ANOMALA (Cooke and Massee) n. comb. Plate xx, figs. 6, 7;
Plate xxi, fig. 9.

Bovista anomala Cke. et Mass., Grev., 1889, xviii, p. 6.—Catastoma anomalum
(Cke. et Mass.) Lloyd, Lyc. Aus., 1905, p. 27.

Peridium up to 2 cm. diam., depressed-globose or pulvinate; exoperidium a thin
brown membrane, flaking away save at the base where persisting as a small
cupulate structure, externally covered with debris;:endoperidium bay brown or
umber, firm, membranous, furfuraceous, dehiscing by a raised, tubular, mammose,
circular stoma which is surrounded by a depressed groove.

Gleba olivaceous, becoming umber, pulverulent; capillitium threads pallid
chestnut brown, of the usual type. Spores globose, 6-8 u diam., stumps only of the
pedicels remaining; epispore pallid chestnut brown, finely and closely verruculose.

Habitat:—Solitary or in small groups on the ground.

Distribution:—Australia.

New South Wales:—*Milson Island, Hawkesbury River, Jan., 1915 (Det. Lloyd
No. 76 as Catastoma anomalum); March, 1916; *Dubbo, June, 1915 (Det. Lloyd
No. 78 as Catastoma anomalum); *Forbes, Aug., 1915 (Det. Lloyd No. 77 as
C. anomalum).

Victoria:—*Ararat, July, 1918, E. J. Semmens.

South Australia: —*Ooldea, Aug., 1922 (2 collections).

Characterized by the thin and membranous exoperidium, finely verruculose
spores and especially by the definite tubular stoma, surrounded by a depressed
groove. The stoma is not always tubular, however, for sometimes it may be
fimbriate and mammose, and in old and weathered specimens almost indefinite.

Sometimes a trace of a gelatinous membrane is present lying on the surface
of the endoperidium. The gleba is frequently tinged with purple in old specimens.

240 GASTEROMYCETES OF AUSTRALASIA, Vii,

6. DISCISEDA AUSTRALIS, n. sp. Plate xxi, figs. 1, 8.

Peridium up to 3 cm. diam., depressed-globose, attached by a small rooting
base; exoperidium thin, umber, fragile, membranous, flaking away irregularly save
at the base; endoperidium furfuraceous, dehiscing by a tubular or mammose
stoma, which is surrounded by a depressed groove.

Gleba olivaceous, pulverulent; capillitium threads pallid chestnut brown, of
the usual type. Spores globose or subglobose, 5-6-5 wu diam., with stumps only of
the pedicels persisting; epispore pallid chestnut brown, closely and finely
verruculose, warts round-topped.

Habitat:—Solitary on the ground.

Distribution:—Australia.

Victoria:—*Ararat, May, 1918, E. J. Semmens. Type, in Herb. Cleland.

South Australia:—*Mt. Pleasant, June, 1925.

The characters of the species are the membranous exoperidium, definite
stoma of the D. anomala type, and the fine, closely grouped, round-topped warts
covering the epispore. These, though readily seen when magnified x 1,000, are
barely perceptible when x 500.

The plant has a general resemblance to the preceding, especially in the
characters of the exoperidium and stoma; but may be separated by the different
spores, larger peridium, and darker colour of the endoperidium.

7. DISCISEDA HYPOGAEA (Cooke and Massee) n. comb. Plate xxii, fig. 2.

Bovista hypogaea Cke. et Mass., Grev., 1891, xx, p. 35.—Catastoma hypogaeum
(Cke. et Mass.) Lloyd, Lyc. Aus., 1905, p. 27.

Peridium up to 2 em. diam., depressed-globose or subglobose; exoperidium
membranous, very thin, pallid ochraceous, fragile, falling away irregularly save
at the base; endoperidium thin, flaccid, papyraceous, olivaceous, furfuraceous,
dehiscing by a minute mammose apical stoma.

Gleba olivaceous, pulverulent; capillitium of the usual type, tinted. Spores
‘globose, 8-9 » diam. (including verrucae), stumps only of the pedicels persisting;
epispore chestnut brown, closely covered with coarse flat-topped echinulae.

Habitat:—Solitary in the ground; hypogaean.

Distribution:—Australia.

New South Wales:—*Narrabri, May, 1919.

The coarse, flat-topped spines of the spores characterize the species. These
are so well marked as to form a definite halo around the spores when these are
viewed in median section; resembling in this particular the spores of certain
species of Geaster. The thin and flaccid nature of the exoperidium is also
characteristic. f

Lloyd (1905, p. 27) records the species as being collected in New Zealand
by R. Brown; the writer has not seen New Zealand specimens.

8. DISscISEDA VERRUCOSA G. H. Cunn. Plate xxi, figs. 2, 6.

Trans. N.Z. Inst., 1926, lvii, p. 205.

Peridium up to 3 cm. diam., depressed-globose; exoperidium brown, tough,
of hyphae and vegetable debris immixed, flaking away irregularly save a small
persistent basal portion; endoperidium thick, tough, membranous, bay brown
or tinged with purple, dehiscing by an irregularly torn, toothed apical stoma.

Gleba purplish, pulverulent; capillitium threads tinted, of the usual type.
Spores globose, 6-8 » diam. (including verrucae), stumps only of the pedicels

BY G. H. CUNNINGHAM. 241

persisting; epispore chestnut brown, covered with coarse hyaline verrucae, often
in the form of finger-like processes.

Habitat:—Solitary or in small groups on the ground.

Distribution:—Australia; New Zealand.

New Zealand:—Milford Track, Otago, Jan., 1920, E. H. Atkinson, Type, in the
herbarium of the writer, No. 887; Waikare, Canterbury, Jan. 1926, G.H.C.

New South Wales:—*Forbes, Aug., 1915.

South Australia:—*Beaumont, May, 1922; *Flinders Range, near Pt.
Augusta, Aug., 1922; *Kinchina, Aug., 1922 (2 collections); *Grange, June, 1925,
Mr. Tee.

The species is characterized by the nature of the spores, these being covered
with coarse, hyaline, finger-like processes. The latter vary in length, however,
in some collections being about half the length of those present in others.

Doubtful and Little Known Species.
DisciseEDA MUELLERI (Berkeley) n. comb.

Bovista Muelleri Berk., Proc. Linn. Soc. Bot., 1878, xvi, p. 171.—Catastoma
Muelleri (Berk.) Lloyd, Lyc. Aus., 1905, p. 27.

Peridium up to 2°5 cm. diam., subglobose, with a short, stout rooting base;
exoperidium soon umber with minute, pale subpersistent warts; endoperidium
firm, rather thick, brown.

Gleba reddish-brown, capillitium threads flaccid, pale, but little branched.
Spores globose, 10-12 uw diam., apedicellate; epispore reddish-umber, coarsely
spinulose.

Habitat:—On the ground.

Distribution:—Australia.

Queensland:—Herberts Creek, Darling Range. Type, in Herb. Kew.

The writer has not seen specimens; the description (slightly modified) being
that given by Cooke in his Handbook (1892). The species is placed in the
doubtful section, for the description of the spores is such that it is not possible
to compare it with the others. According to Lloyd this has the roughest spores
of any species included in the genus. In the description the gleba is said to
be reddish-brown, and in fact this is the only character by which it can be said
to differ from most others placed in the genus; but as has been shown, glebal
colour is not of specific value.

A single broken specimen of some Disciseda in the possession of Dr. Cleland
has a similar reddish-brown gleba, but the spores are much smaller than the
measurements given for D. Muelleri and are angular and tuberculate-striate.
The specimen is too broken to allow of its being described.

Lloyd (Myc. Notes, 1917, p. 631) describes a species under the name of
Catastoma magnum, stated to have been collected in Canterbury. He considered
it differed from D. anomala in the larger size, thick, leathery exoperidium and in
the apedicellate, strongly roughened spores, but claimed it was but a form of
D. anomala. From his description it is impossible to say to what species it should
be referred. The same “species” has been recorded from South Africa by
Verwoerd (1925).

ApsTtomMa G. H. Cunn.
Trans. N.Z. Inst., 1926, lvii, p. 206.
Peridium subglobose, of two layers; a thick, fragile exoperidium composed
of hyphae immixed with sand particles, breaking away irregularly; and a

249 GASTEROMYCETES OF AUSTRALASIA, Vii,

papyraceous or membranous, coloured endoperidium which dehisces by irregular
rupture, a stoma being absent.

Gleba of spores and capillitium, compact or pulverulent at maturity;
capillitium threads short, occasionally branched, smooth, continuous, coloured.
Spores apedicellate, globose, reticulate, coloured.

Habitat:—Solitary or in small groups in the ground; hypogaean.

Distribution:—Australia; New Zealand.

The genus is separated from the preceding because of the absence of a stoma,
dehiscence being effected by the irregular breaking away of the peridium; in
other characters, as habit, structure, capillitium, etc. it is practically identical.
The spores, however, are characteristic in the two species described below in
that they are reticulate, a condition not Known to occur with any species of
Disciseda.

1. ABSTOMA PURPUREUM (Lloyd) G. H. Cunn. Plate xxi, fig. 3; Plate xxii, fig. 4.

Trans. N.Z. Inst., 1926, lvii, p. 206.—Catastoma purpurea Lloyd, Myc. Notes.
1922, p. 1120.

Peridium up to 4 cm. diam., subglobose or depressed-globose; exoperidium a
sand-case, up to 3 mm. thick, firm, brittle, dull purple, breaking away irregularly
and completely; endoperidium thin, papyraceous, fragile, dark purple, almost
black, free from the exoperidium save at the base where attached (but not
organically); dehiscing by irregular rupture.

Gleba purple, firm and compact; capillitium threads deep chestnut brown,
thick walled. Spores globose or shortly elliptical, 8-16-5 « diam., apedicellate;
epispore deep chestnut brown, closely and finely reticulate.

Habitat:—In sand among dunes; hypogaean.

Distribution: —New Zealand.

Weraroa, Wellington, Nov., 1919, S. A. Cunningham, HE. H. Atkinson, G.H.C.
Type collections, and type of the genus, in the herbarium of the writer, No. 1783.

The species is characterized by the purple colour of the whole plant, the
thick sand-case exoperidium, fragile, non-stomate endoperidium, and deeply
coloured, reticulate spores.

The habit is peculiar. The plant grows buried to a depth of several
centimetres, 7-10 or more, in sand at Weraroa beach. When wind erosion occurs,
as is frequent in this area of shifting dunes, the sand is blown for some
distance, large amphitheatres being formed, and the globose plants are carried by
the wind often for one kilometre or more from their place of origin. They are
rolled along the surface of the dune slopes and finally arrested by debris of the
nature of branches of trees and stumps half buried at the tops of the long dune
slopes. Here they come to rest and the peridium is gradually removed by friction
of the sand particles.

2. ABSTOMA RETICULATUM, n. sp. Plate xxii, fig. 3.

Peridium up to 3 cm. diam., depressed-globose; exoperidium of the nature
of a fugacious sand-case; endoperidium umber or sepia brown, tough, membranous,
smooth, dehiscing by irregular rupture.

Gleba olivaceous, pulverulent; capillitium threads tinted, thin walled. Spores

globose, 8-12 w diam., apedicellate; epispore coarsely and strongly reticulate,
chestnut brown.

Habitat:—Solitary on the ground; hypogaean?

BY G. H. CUNNINGHAM. 243

Distribution:—Australia.

New South Wales:—*Forbes, Aug., 1915, J. B. Cleland. Type, in Herb. Cleland.

The olivaceous colour of the gleba, tough and membranous endoperidium, and
coarsely reticulate spores separate the species from the preceding. The
exoperidium has almost completely disappeared from this collection.

Literature Cited.

CooxE, M. C., 1892.—Handbook of Australian Fungi, 457 pp., London.
CUNNINGHAM, G. H., 1926.—Lycoperdaceae of New Zealand. Trans. N.Z. Inst., lvii,
p. 187.
CZERNAIAIEV, B. M., 1845.—Nouveaux cryptogames de 1l’Ucraine. Bull. Soc. Imp. de
Moscow, xvili, p. 153. ;
Ho.utos, L., 1902.—A Disciseda Czern. genusz fajai. Noevenytani Koezlemenyek, 1. p. 105
, 1902.—Gasterom. vonatk. helyesb. Termeszetr. Fuwez., xxv, p. 100.
, 1903.—Die arten der Gattung Disciseda Czern. Hedwigia, xlii, p. 20.
Luoyp, C. G., 1901 et seq.—Mycological Notes, Cincinnati.
, 1905.—The Lycoperdaceae of Australia, New Zealand and Neighboring Islands.
44 pp., Cincinnati.
Morean, A. P., 1892.—North American Fungi, V. Journ. Cincinnati Soc. Natural History,
xiv, p. 141.
VERWOERD, LEN, 1925.—Suid-Afrikaanse Lycoperdaceae en Nidulariaceae. Annale van
die Uniwersiteit van Stellenbosch, Jaargang 3, Reeks A, No. 1, p. 1.

Index.
Synonyms are in italics.
Abstoma G. H. Cunn. Catastoma Morg. (continued)
purpureum (Lloyd) G. H. Cunn. Muelleri (Berk.) Lloyd
reticulatum G. H. Cunn. pedicellata Morg.
Bovista anomala Cke. et Mass. purpurea Lloyd
argillacea Pat. . subterranea (Peck) Morg.
candida Schw. Disciseda Czern.
cervina Berk. Key to species (p. 236)
circumscissa Berk. et Curt. anomala (Cke. et Mass) G. H. Cunn.
debreciensis (Hazsl.) de Toni australis G. H. Cunn.
hyalothrix Cke. et Mass. candida (Berk.) Lloyd
hypogaea Cke. et Mass. cervina (Berk.) G. H. Cunn.
Muelleri Berk. circumscissa (Berk. et Curt.) Hollos
subterranea Peck debreciensis (Hazsl.) Hollos
Catastoma Morg. hyalothrix (Cke. et Mass.) Hollos
anomalum (CKe. et Mass.) Lloyd hypogaea (CKe. et Mass.) G. H. Cunn.
circumscissa (Berk. et Curt.) Morg. Muelleri (Berk.) G. H. Cunn.
debreciense (Hazsl.) Hollos pedicellata (Morg.) Hollos
hyalothrix (Cke. et Mass.) Lloyd verrucosa G. H. Cunn.
hypogaeum (Cke. et Mass.) Lloyd Globaria debreciensis Hazsl.

magnum Lloyd

EXPLANATION OF PLATES XX-XXII.

it Plate xx.

Fig. 1. Disciseda pedicellata, natural size. The upper plant is reversed to show the
persistent basal portion of the exoperidium.

Fig. 2. D. pedicellata, x 2. Showing the plane stoma and the smooth endoperidium.

Fig. 3. D. candida, natural size. Note the large mammose stoma.

Fig. 4. D. candida, x 3. Showing the furfuraceous endoperidium, persistent basal
portion of the exoperidium and the fimbriate mammose stoma.

Fig. 5. D. cervina, natural size. Note the mammose stoma and persistent basal
portion of the exoperidium.

Fig. 6. D. anomala, natural size.

Fig. 7. D. anomala, x 3:5. Note the tubular mammose stoma surrounded by a depressed

ring.

244 GASTEROMYCETES OF AUSTRALASIA, Vii.

Plate xxi.

Fig. 1. D. australis, natural size.
Fig. 2. D. verrucosa, natural size.
Fig. 3. Abstoma purpureum, natural size.
Fig. 4. Disciseda pedicellata, spores, x 1200.
Fig. 5. D. hyalothrix, spores, x 1200.
Fig. 6. D. verrucosa, spores, x 1200.
Fig. 7. D. candida, spores, x 1200.
Fig. 8. D. australis, spores, x 1200.
Fig. 9. D. anomala, spores, x 1200.

Plate xxii.
Fig. 1. Disciseda cervina, spores, x 1200.
Fig. 2. D. hypogaea, spores, x 1200.
Fig. 3. Abstoma reticulatum, spores, x 1200.
Fig. 4. A. purpurewm, spores, x 1200.

All photographs (except Plate xxi, fig. 3, by H. Drake) have been taken by the
writer. The spore drawings have been made with the aid of a camera lucida; x 1200.

THE GASTEROMYCETES OF AUSTRALASIA.
viii. THE GENUS MYCENASTRUM.
(Plate xxii, figs. 5-8.)

This genus contains a single species, which has been collected in practically
every country in the world, though it is most abundant in sandy regions.

The peridium ranges in size from 2 cm. to 20 cm. or more, and may be globose,
obovate or pyriform. It consists of the usual two tissues, exoperidium and
endoperidium.

The exoperidium is thin, floccose and usually smooth externally, but forms
occur in which the flocci are aggregated into definite scales; at maturity it
usually falls away completely, exposing the endoperidium. This latter tissue is
unlike that of the other genera of this sub-family in that it is thick, 2-5 mm.,
and indurated. In young plants it may appear almost cartilaginous, but in
mature plants it is hard and coriaceous, or even corky.

The peridium may dehisce in a stellate manner, or by the falling away in
irregular flakes from the apical portion.

The gleba consists of spores and capillitium, and is decidedly pulverulent at
maturity. The capillitium threads are peculiar and characteristic. Each consists
of a short main stem which may be simple or short-branched, and is covered
with short, acuminately pointed spines. These threads differ considerably, even
in the same plant, in the degree of spininess, shape and size. Mixed with the
capillitium threads are usually present numerous long and unbranched septate
hyphae of a type similar to those forming the peridium.

MycenASTRUM Desvaux.

Ann. Sci. Nat., 2, 1842, xvii, p. 143.—Pachyderma Schulz., Verh. Zoo. Bot. Ges.
Wien., 1875, xxv, p. 79.

Peridium globose, obovate or pyriform, of two layers; a thin, floccose
exoperidium; and a thick, indurated, persistent endoperidium; dehiscing in a
stellate manner, or by the irregular rupture of the apical portion; sterile base
absent.

Gleba olivaceous, becoming umber, pulverulent; capillitium threads very
abundant, of numerous short hyphae, continuous, short-branched or simple,
branches beset with stout, spinous processes. Spores globose or elliptical, coloured,
coarsely echinulate.

Habitat:—Solitary, in small groups or caespitose on the ground; epigaean.

Distribution:—Hurope; North America; Asia; India; Africa; Australia; New
Zealand.

The genus is characterized by the nature of the capillitium, the hard and
coriaceous endoperidium and the method of dehiscence.

MYCENASTRUM coRIUM (Guersent) Desvaux. Plate xxii, figs. 5-8.

Ann. Sci. Nat., 1842, xvii, p. 147.—Lycoperdon corium Guers., in DC. FI. Fr.,
suppl. 2, 1815, p. 598. Bovista suberosa Fr., Syst. Myc., iii, 1829, p. 26. Scleroderma

246 GASTEROMYCETES OF AUSTRALASIA, Viii.

corium Grev., in Duby Bot. Gall., ii, 18380, p. 892. Mycenastrum phaeotrichum
Berk. in Hook. Lond. Jour. Bot., ii, 1843, p. 418. M. chilense Mont., Ann. Sci. Nat.,
ser. 2, xx, 1843, p. 375. Hndoneuron suberosum Czern., Bull. Soc. Imp. Mosc.,
xviii, 1845, p. 152. Mycenastrum leptodermeum Dur., Fl. Alg., 1849, p. 386.
M. radicatum Dur., l.c., p. 387. M. Beccarii Pass., Nuov. Giorn., vii, 1875, p. 183.
Pachyderma Strossmayeri Schulz., Verh. Zoo. Bot. Ges. Wien, xxv, 1875, p. 79.
Mycenastrum clausum Schulz., I[bid., xxvii, 1877, p. 114. Bovista spinulosa Peck,
Bot. Gaz., iii, 1879, p. 170. Mycenastrum spinulosum Peck, Thirty-Third Rept. for
1879, 1883, p. 15. M. olivaceum CKe. et Mass., Grev., xvi, 1887, p. 33. Scleroderma
phaeotrichum (Berk.), de Toni, in Sacc. Syll. Fung. vii, 1888, p. 139. S. chilense
(Mont.) de Toni, l.c. S. spinulosum (Peck) de Toni, l.c. S. radicatum (Dur.)
de Toni, Z.c. S. leptodermeum (D. et M.) de Toni, l.c. S. Beccarii (Pass.) de Toni,
lc. S. olivaceum (Cke. et Mass.) de Toni, 1.c.

Peridium globose, subglobose, obovate or pyriform, up to 20 cm. diam.;
exoperidium tomentose, fugacious, greyish; endoperidium thick, 2-5 mm., smooth,
polished, at first greyish. becoming bay brown, dehiscing in a stellate manner,
or by the irregular falling away of the apical portion.

Gleba olivaceous, becoming umber, pulverulent; capillitium threads of the
usual type. Spores globose or shortly elliptical, 11-13 mw diam., apedicellate;
epispore chestnut brown, densely echinulate, reticulate, wall 2 mw thick.

Habitat:—Solitary, in groups or caespitose on the ground.

Distribution:—Same as that of the genus.

Queensland:—*Barron Falls, Kuranda, Sept., 1917.

South Australia:—*Glen Osmond, Dec., 1920; *Adelaide, Oct., 1922; Feb., 1921;
*Hncounter Bay, Jan., 1924; *Pt. Elliot, Aug., 1918, D. I. Cleland.

Western Australia:—*Kalgoorlie, June, 1916.

New South Wales:—*Dungog, Nov., 1916; *Moore Park, Sydney. Dec., 1917;
*Mudgee, Dec., 1917; *Cowra Experimental Farm, Dec., 1917; *Coolamon, May,
1918; *Narrabri, June, 1919.

New Zealand:—Castlepoint, Wairarapa, Jan., 1923, Miss Edwin; Spring Creek,
Blenheim, Jan., 1925, W. D. Reid; Levels, Canterbury, Jan., 1927, G.H.C.

The species varies greatly in the size of the peridium, degree of spininess
of the capillitium, roughness of the spores, and colour of the gleba. Usually
if plants are collected and dried when young the gleba appears olivaceous; if
dried when mature it may be umber or even purple. ‘

Caespitose plants are frequent, at least in Australia and New Zealand. Plants
break away from the point of attachment at maturity and are then often carried
for considerable distances by wind, being rolled along the ground, and in this
manner the spores are widely dispersed.

EXPLANATION OF PLATE XXII.

fe
OQ
on

Mycenastrum corium. x %. Showing the thick endoperidium and the stellate

method of dehiscence. (Photo. H. Drake.)

Fig. 6. M. corium. x 4. Showing the flaking exoperidium. (Photo. by writer.)

Fig. 7. Capillitium. x 125. Showing the spinose nature of the threads; air bubbles
in the lumen of the threads show dark in this photo. (Photomicrograph by
writer.)

Fig. 8. Spores. x 1200. Drawn from a lactic acid mount with the aid of a camera

lucida; New Zealand form. 'The spines are somewhat exaggerated.

* Collections in the herbarium of Dr. J. B. Cleland, Adelaide University.

THE GASTEROMYCETES OF AUSTRALASIA.
ix. Krys TO THE GENERA AND SPECIES OF THE LYCOPERDACEAE.

By G. H. CUNNINGHAM, Mycologist, Department of Agriculture,
Wellington, N.Z.

[Read 27th July, 1927.]

The writer would limit the Lycoperdaceae to those genera possessing a non-
stipitate, two- or four-layered peridium, copious capillitium, and long-sterigmate,
4- or 8-spored basidia. The limits of the family have been much greater in the
past, as the following classifications will show, but as our knowledge of the
development and structure of the various genera have broadened, so the limits
of the family have been narrowed.

Details of the various classifications which have been proposed are of little
other than historical interest today, to the student of this group, but a few are
given here to indicate the diversities of opinion held by different authors, as to
the position and affinities of the genera included in the Lycoperdaceae.

Commencing with Persoon’s Synopsis Methodica Fungorum, 1801 (the starting-
point of Modern Nomenclature in the Gasteromycetes), we find he divides the
Fungi as under (details of the Gasteromycetes only are given) :—

1Class I. ANGIOCARPI: Fungi closed, or bearing very numerous spores internally.

Order 1. Sclerocarpi.
2. Sarcocarpi. Fungi fleshy stuffed. Sphaerobolus, Thelebolus.
8. Dermatocarpi. Fungi membranous, tough, internally powdery.
+Trichospermi. Seed powder mixed with threads. 13. Batarrea.
14. Geastrum2 15. Bovista. 16. Tulostoma. 17. Lycoperdon.
18. Scleroderma (Genera 19-27 are now placed in a separate
group, the Myxomycetes).

+7Gymnospermi. Seed powder without threads.
777Sarcospermi. Fructifications abundant, fleshy. 38. Cyathus.

Class II. GymMNnocarPi: Fungi fleshy. Seeds (few) borne in an open receptacle.
Order 4. lLytothecii. Fertile membrane at length mucilaginous. 39. Clathrus.
40. Phallus.
5. Hymenothecii (Hymenomycetes).
6. Naematothecii (Hyphomycetes).

As this and other classifications of that period are not in use today, the many
others may be omitted, and attention paid only to those which have been in general
use in recent years. Perhaps the best known of these is that followed in Saccardo’s
Sylloge Fungorum, Vol. 7, 1888, where the Lycoperdaceae is arranged as under

1Thanks are due to Miss Wakefield, Kew Herbarium, for a translation of Persoon’s
classification. Unfortunately this work is not procurable in New Zealand.
2Genera included by the writer in the Lycoperdaceae are italicized; those excluded
by him, or not present in Australasia, are in ordinary type. of
F

248 GASTEROMYCETES OF AUSTRALASIA, ix,

(only the arrangement of this family in the various classifications will be
considered henceforth) :—

Family III. Lycoperdaceae.
Sub-family 1. Podaxineae.

2. Diplodermeae.
Tylostoma, Queletia, Battarrea, Husseya, Mitremyces,
Geaster, Diplocystis, Diploderma, Trichaster, Broomeia,
Coilomyces.

3. Lycoperdeae.
Lanopila, Eriosphaeria, Bovista, Calvatia, Lycoperdon.

4. Sclerodermeae.
(Mycenastrum is placed here as a synonym of Scleroderma).

Ed. Fischer in Engler and Prantl’s Natuerliche Pflanzenfamilien, I, 1**, 1900,
arranged the family as under:—
Order 3. lLycoperdineae.
Family I. Lycoperdaceae.

Lycoperdon, Globaria, Catastoma (= Disciseda), SBovistella, Bovista,
Mycenastrum, Geaster.
Lloyd’s classification (Genera of the Gasteromycetes, 1902) is as follows:—
Family 4. Lycoperdaceae.
Tribe 1. Tylostomeae.
2. Podaxineae.
3. Sclerodermeae.
4. Lycoperdeae.
Tribal Alliance i. Geastreae.
Geaster.
ii. Bovisteae.
Bovista, Mycenastrum, Catastoma (Ee
Disciseda).
ili. Lycoperdeae.
Bovistella, Lycoperdon, Calvatia.
iv. Anomalae.
Gallacea, Castoreum, Arachnion, Meso-
phellia, Mitremyces.
Hollos (Gasteromycetes Ungarns, 1904), follows the arrangement :—
Family 3. Lycoperdaceae. F
Battarea, Tylostoma, Myriostoma, Geaster, Astraeus, Calvatia,
Lycoperdon, Disciseda, Bovista, Mycenastrum.
Carleton Rea (British Basidiomycetae, 1922) arranges the family as under:—
Sub-order 3. Lycoperdineae.
Family 1. Lycoperdaceae.
Lycoperdon, Bovistella, Bovista, Myriostoma, Geaster.

From a consideration of these classifications we find that Persoon included
Battarrea, Tulostoma, Scleroderma and the Myxomycetes in the family; that
Saccardo placed Geaster in a separate sub-family, Mycenastrum as a synonym of
Scleroderma, and included with Lycoperdon, etc. such genera as Lanopila and
HBriosphaeria, while Tulostoma, Mitremyces (= Calostoma), etc. are placed in the
sub-family Diplodermeae; that Fischer placed the genera later considered by the
writer all under the Lycoperdaceae; that Hollos followed the older school in
considering under the Lycoperdaceae such genera as Battarrea, Tulastoma, ete. ;
and that Rea’s arrangement was similar to that of Fischer.

It is out of place to consider in detail the reasons why such genera as
Tulostoma, Battarrea, Scleroderma, etc. do not belong to this family; these
reasons will be given in subsequent papers dealing with these genera. As has
been shown, the writer limits the family to those genera possessing a copious
eapillitium (which excludes Scleroderma); a non-stipitate peridium (which

BY G. H. CUNNINGHAM. 249

excludes Tulostoma and Battarrea); 4- or 8-spored, sterigmate basidia (which
excludes such genera as Diploderma and its allies).

All the genera considered as belonging to the family (Abstoma, Bovista,
Bovistella, Calvatia, Disciseda, Geaster, Lycoperdon, Mycenastrum (extra-
Australasian genera are not considered here) possess a two- or (in Geaster) four-
layered peridium; the gleba is composed of spores immixed with copious
capillitium threads; no stipe is present in any genus; the basidia are all long-
sterigmate, and bear apically 4-8 unicellular spores. The development has been
worked out for all genera by the writer, and is in most (save Geaster) found
to be similar to that of Lycoperdon, save in minor details, as the formation of the
capillitium threads.

STRUCTURAL SIMILARITIES AND DIFFERENCES IN THE GENERA OF THE LYCOPERDACEAE.

Tribe Lycoperdeae.

In the mature plant a two-layered peridium is always present, and the plants
are attached to the substratum by a rooting base (Bovistella and Lycoperdon)
which holds them to the place of origin even when old and withered; or they may
be attached by a small rooting strand or mycelial cord and at maturity break
away and be carried by the wind for some distance (Abstoma, Bovista, Disciseda
and Mycenastrum).

The outer layer of the peridium is termed the exoperidium, and is usually
pseudoparenchymatous, but may be membranous, and is often composed of warts
and spines. This structure is often quite thin and flakes away in irregular patches.
In Abstoma and Disciseda this membrane is frequently a thick (1-3 mm.) sand-
case, consisting of loosely woven hyphae with which are immixed sand grains or
particles of vegetable matter. The whole structure though firm is quite brittle,
and falls away in irregular fragments. The exoperidium is one of the most
variable tissues, ranging in the genus Lycoperdon from a smooth membrane,
through furfuraceous to verrucose and even echinulate. The spines or verrucae
seldom fall away completely but usually partly persist, especially towards the
base. These structures are usually considered valuable aids in diagnosis, but
should be used with caution, as in certain species individual plants in the same
collection may possess either furfuraceous, verrucose or even echinulate processes
to the exoperidium. In those species which are conspicuously spinose, the spines
are usually surrounded at their bases with rings of small granules; consequently
when the spines fall, the granules persist, giving to the exposed endoperidium a
curiously reticulated appearance (LL. compactum).

The inner tissue, the endoperidium, is in most genera thick and membranous
or papyraceous, but in Mycenastrum is thick (2-5 mm.), corky, coriaceous and
pseudoparenchymatous, and in growing plants often cartilaginous.

The peridium encloses the gleba, which at maturity consists of capillitium
threads and spores. The threads serve aS most useful generic characters. In
many species of the genera Bovistella and Lycoperdon is present the so-called
columella, a cylindrical or elliptical structure of sterile tissue traversing the
lower portion of the gleba and attached to the base of the plant. In certain species
of the genera Bovistella, Calvatia and Lycoperdon a sterile base is present. This
consists of a usually cellular tissue occupying the lower portion of the gleba, and
is most frequent in those plants with a well-developed, stem-like base. In two
species (Lycoperdon polymorphum and Calvatia candida) the sterile base is not
cellular but compact, and consequently frequently difficult to differentiate from

250 GASTEROMYCETES OF AUSTRALASIA, ix,

the gleba. In certain species of Calvatia and Lycoperdon (Calvatia lilacina,
C. caelata, Lycoperdon depressum) the sterile base is separated from the gleba by
a definite diaphragm.

The spores are continuous, globose or shortly elliptical, and possess rough
or smooth epispores. Their shape, size and degree of roughness are useful
specific characters. They may have long and slender sterigmata (often of irregular
size on the same basidium) which in turn are borne apically on small cylindrical
or clavate basidia. When the sterigmata with the attached spores break away
from the basidia the latter are said to be pedicellate; when stumps only of the
sterigmata remain attached to the spores the latter are said to be apiculate (though
the use of this term is not strictly accurate, it being used generally in the sense
of an apical point or apicula, the pedicel being basal); or should the spores become
completely detached from the sterigmata they are said to be apedicellate. 'The
basidia in all members of this tribe are regularly tetrasporous.

Most of the genera show a general resemblance, so close in fact that difficulty
is sometimes experienced in separating species of Bovistella from Lycoperdon or
Bovista. In the writer’s experience separation is best effected by means of the
capillitium threads, and method of dehiscence. In Calvatia and Lycoperdon the
threads are very long, even throughout their length, sparingly septate or
continuous. They are free at one end, and usually attached by the other to the
inner wall of the peridium, or to the columella and diaphragm or sterile base
should these be present. In Lycoperdon three types of capillitium threads are
present; hyaline, septate, simple threads; coloured, continuous, simple or sparingly
branched threads; and coloured, freely branched threads. These last approach in
appearance those of Bovista and Bovistella.

Calwatia is separated from Lycoperdon chiefly by the method of dehiscence.
In the former genus the upper portion of the peridium flakes away in pieces, when
at maturity the whole may have disappeared, the gleba then being directly exposed.
In Lycoperdon dehiscence is effected by a definite apical stoma. In two species
(L. depressum and L. Wrightii) dehiscence is first effected by means of the apical
stoma, but as the plant ages, irregular flaking away of the whole of the apex
of the peridium occurs; consequently difficulty might be experienced in placing
plants of these species, according to whether they were collected when mature or
immature. In such cases the peculiar hyaline, septate capillitium serves to
separate them.

In Abstoma and Disciseda the capillitium threads are very short, unbranched
and usually continuous; and are so characteristic as usually to place any species
in these two genera. Immature plants possess capillitium approaching the
Lycoperdon or Calvatia type. In Disciseda dehiscence is effected by a stoma which
in most species is apical but in one or two species is said to be basal. In Abstoma
no stoma is present, the spores being liberated by irregular rupture of the
endoperidium.

Bovista and Bovistella both possess capillitium threads which are peculiar
and characteristic. Each thread is free, and has a short, distinct, thick main-
stem, with numerous short, dichotomous, acuminate branches. The threads are
penetrated to the lumen by peculiar conic pits (absent in Bovista purpurea).
Dehiscence is effected by means of an apical stoma, as in Lycoperdon. The two
genera are separated from one another upon habit, Bovista being a genus in which
plants break away at maturity from the place of growth, Bovistella persisting by
means of a rooting base as does Lycoperdon. Herbarium specimens may usually

BY G. H. CUNNINGHAM. 251

be differentiated by the presence or absence of this rooting base. This in turn is
usually indicated by the presence of a sterile base, so that by many workers,
Bovistella has come to be recognized as a Bovista with a sterile base. In the
opinion of the writer, however, this is not a sufficiently valid distinction upon
which to base a genus.

Mycenastrum is characterized both by the capillitium threads and method of
dehiscence. The threads are short, free, short-branched and covered with numerous
spinous processes. Dehiscence is effected by the stellate splitting of the apex, but
in certain forms, even in the same collection, it may be effected by the irregular
breaking away of the apical portion.

Tribe Geastreae.
This is represented in Australasia by the single genus Geaster.

The structure of species of this genus differs in that no less than four well-
defined tissues are present: (1) An outer mycelial layer consisting of a palisade
of stout fibres; (2) a central fibrillose layer of interwoven hyphae with long axes
predominantly radial; (3) an inner fleshy layer, definitely pseudoparenchymatous
in structure (these three tissues comprise the exoperidium); and (4) the
endoperidium.

The exoperidium splits into a variable number of rays, which may be
expanded, revolute or involute. Dehiscence is effected by the exoperidium splitting
from the apex downwards into several stellate lobes, exposing the endoperidium,
which in turn dehisces by an apical stoma. The endoperidium is usually
membranous or papyraceous, and may be carried on a short pedicel, or be sessile.
The stoma may be a poorly defined aperture, scarcely discernible from the
endoperidium—when it is said to be naked; or it may be enclosed within a definite
peristome, when it is said to be peristomate. The peristome may be externally
fluted, when it is termed plicate; or silky-fibrillose, when it is said to be fibrillose.

The capillitium threads are continuous, simple (save in Geaster velutinus
where they are sometimes branched at their apices), thick-walled, acuminately
pointed and dark-coloured. The basidia are usually 4-spored, but 8-spored forms
are not uncommon. A columella is present in certain species, but is noticeable
usually only in immature specimens immediately before dehiscence.

The spores are usually globose, usually some shade of brown, and mostly
verrucose or verrucose-echinulate.

In the family Lycoperdaceae there are 57 Australasian species, distributed
among the following eight genera: Abstoma 2, Bovista 2, Bovistella 3, Calvatia 4,
Disciseda 8, Geaster 23, Lycoperdon 14, and Mycenastrum 1.

Their distribution is interesting, as the accompanying table shows.

Table of Distribution of the Species.

Species. Distribution.
Abstoma purpureum sel wiyeig asi sp palpetrare hs New Zealand
Abstoma reticulatum OME Reichs noodle. Wacom Australia
Bovista brunnea PL Meets! Bees hh Woelisl i sigeeaie ened New Zealand, Australia
IDOWISHO (OUMOUFAG. og co 60, nb 65. OC New Zealan
Bovistella vovistoides 5 “22 ae. New Zealand, Australia, India |
TEHRAN eS FOURS oo oo on ao 95. oD Australia ie
Bovistella verrucosa see) any Mi ISR erence anG Australia te

G \@

252

GASTEROMYCETES OF AUSTRALASIA, ix,

Species.

Calvatia caelata

Calvatia candida
Calvatia gigantea

Calvatia lilacina

Disciseda anomala
Disciseda australis
Disciseda candida

Disciseda cervina ..
Disciseda hyalothrizx
Disciseda hypogaea
Disciseda pedicellata
Disciseda verrucosa

Geaster
Geaster
Geaster
Geaster
Geaster
Geaster
Geaster
Geaster
Geaster
Geaster
Geaster

arenarius
australis
Berkeleyi
Bryantit
campester
Clelandit
Drummondii
ellipticis~
fenestriatus
jimbriatus
floriformis

Geaster Harioti
Geaster limbatus

Geaster minus

Geaster mirabilis

Geaster pectinatus

Geaster plicatus
Geaster saccatus

Geaster simulans
Geaster Smithii ‘
Geaster subiculosus
Geaster triplex

Geaster velutinus

Lycoperdon
Lycoperdon
Lycoperdon
Lycoperdon
Lycoperdon
Lycoperdon
Lycoperdon

Lycoperdon
Lycoperdon
Lycoperdon
Lycoperdon
Lycoperdon

Lycoperdon
Lycoperdon

asperum
compactum
depressum
glabrescens
Gunnii
nitidum
perlatum

piriforme
polymorphum
pusillum
scabrum
spadiceum

stellatum
subincarnatum

Mycenastrum corium

Distribution.

New Zealand, Australia. North America,
Europe, Asia, Africa

Australia, Europe

New Zealand, Australia, North America,
Europe

New Zealand, Australia, North America,
Europe, Africa

Australia

Australia

New_ Zealand, Australia, North America,
Europe, South America

Australia, North America, Europe

Australia

New Zealand, Australia

Australia, North America, Africa

New Zealand, Australia

Australia, North America

Australia

Australia, Europe

Australia, North America, Europe

Australia, North America, Europe

Australia

Australia

Australia

Australia, North America, Europe, Africa

Australia, North America, Europe

New Zealand, Australia, North America,
Europe, Africa

Australia, Europe, Africa, South America

New Zealand, Australia, North America,

Europe, Africa

New Zealand, Australia, North America,
Europe, Asia

Australia, North America, Asia, South
America

New Zealand, Australia, North America,
Europe

New Zealand, Australia, Africa, India

Australia, North America, Europe, South
America

Australia

Australia, North America

Australia

New Zealand, Australia, North America,
Europe, South America

New Zealand, Australia, North America,
Africa, South America

Australia, Africa, South America

New Zealand

New Zealand, Australia, Europe, Africa

New Zealand, Australia

Australia

Australia

New Zealand, Australia, North America,
Europe, Africa, South America, India

New Zealand, Australia, North America,
Europe, South America, Asia

New Zealand, Australia, North America,
Europe, Africa

New Zealand, Australia, North America,
Europe, Africa, Asia, India

New Zealand, Australia

New Zealand, Australia, Europe

Australia

Australia, North America

New Zealand, Australia, North America,
Europe, Africa, Asia, India

This table shows that of these 57 species, 27 occur in New Zealand; 54 in
Australia; 24 in Europe; 14 in Africa; 5 in India; 26 in North America; 9 in
South America; 6 in Asia; 23 being confined to this biological region.

In the genus Geaster (and the related extra-Australian genus Myriostoma) are
certain structural differences that separate it from the other genera placed in

BY G. H. CUNNINGHAM. 253

the family. Consequently the family may conveniently be divided into the two

following tribes :—

Tribe Lycoperdeae: Peridium two-layered, dehiscing by an apical stoma or by the
gradual falling away of the upper portions of the peridium; capillitium attached
or free, simple or freely branched; basidia sterigmate, 4-spored.

Tribe Geastreae: Peridium four-layered, dehiscing by the exoperidium splitting from
the apex downwards in a stellate manner exposing the endoperidium, which
in turn dehisces by means of an apical stoma; capillitium attached, simple;
basidia sterigmate 4-8 spored.

The genera may be arranged as under :—

Key to the Genera.
Tribe Lycoperdeae: Peridium 2-layered; basidia 4-spored.

Capillitium threads attached to the endoperidium and central columella, when present.
Peridium dehiscing by irregular apical rupture ......... 1. Calvatia.
Peridium dehiscing by a definite apical stoma .......... 2. Lycoperdon.

Capillitium threads free within the endoperidium.

Threads simple, or short-branched.
Threads short, simple, smooth.

Stoma presents) vapicall or balsalleoria cvcroictete clei cnetereie 5. Disciseda.
Stona, APSE ty. cpr easerer sieMowen oie ielocue aver eckev este ven einer each anetten ee 6. Abstoma.
Mhreads) shorts simples sSpiInOSe erste cc ciciaicieitetelel lee 7. Mycenastrum.

Threads freely branched, of a central main stem, and short, dichotomous,
acuminate branches.

Plants with a strong rooting base .................- 3. Bovistella.
Plants without a definite rooting base ............... 4, Bovista.
Tribe Geastreae: Peridium 4-layered; basidia 4-8 spored.
Peridium with a Solitanywaplcalsstomamenerie ote eine eke 8. Geaster.

In order to bring all species of the family together, the following keys to the
species of each genus have been given.

1. CALVATIA Fries.
(For emended descriptions, see Proc. LINN. Soc. N.S.W., 1926, li, p. 363.)

Diaphragm present.

SPOLresuSmMOOthy oko ccs ial Oe Aa tone Rae Me eae ls C. caelata (Bull.) Morg.

SDPOLESMVELLUCOSE | pa. ac die ssncre ce fie eter chao emeped nieeeyeusteb eae oe aGhe = C. lilacina (B. et M.) Lloyd
Diaphragm absent.

Plants large, exterior smooth and leathery .... C. gigantea (Pers.) G. H. Cunn.

Plants small, exoperidium furfuraceous .......... C. candida (Rostk.) Hollos

2. iyCoPERDON Tourn. ;
(Descriptions, Proc. LINN. Soc. N.S.W., 1926, li, p. 627.)

Spores without distinct pedicels.
Capillitium hyaline, freely septate.

Diaphragm present PER Agnic o iota, loid/o 0106.0 O MIO IAI AOE ORGS LL. depressum Bon.
Diaphragm absent.
Exoperidium with minute depressions ........ L. subincarnatum Peck

Exoperidium reticulate with persistent granules .....................
L. compactum G. H. Cunn.

Capillitium coloured, usually continuous.
Capillitium continuous or sparingly branched.
Sterile base of large cells, 2 mm. or more.

Exoperidium of minute connivent spines .... L. piriforme Schaeff.
Exoperidium of pointed verrucae .............. L. perlatum Pers.
Sterile base of minute cells, 1 mm. or less.
Exoperidium of large cruciate spines .... L. stellatwm Cke. et Mass.
MXOperidiumprLurtura couse precise L. nitidum Lloyd
Capillitium freely branched.
Sterile: (base) cellular sc is fais tae sideipoaklie liao ise os L. spadiceum Pers.
Sterilenbase compacts seacaverneacwelexshovonshoroncrekonclinicieieieke L. polymorphum Vitt.

Sterilesibaise: -aAbsenty Hosters srs s Aone maior wevalo euetcin mia e L. pusillum Pers.

254 GASTEROMYCETES OF AUSTRALASIA, ix,

Spores long pedicellate.
Sterile base well developed.

HM XOperidilumeturturaceousmec aero L. glabrescens Berk.

Exoperidium of cruciate spines .......... LI. scabrum (Uloyd) G. H. Cunn.
Sterile base scanty or absent.

Exoperidium of pallid cruciate spines ......... LL. asperum (Lev.) de Toni.

Exoperidium furfuraceous or tomentose .................. LL. Gunnit Berk.

3. BOVISTELLA Morg.

(Descriptions, Proc. Linn. Soc. N.S.W., 1925, 1, p. 370.)
Spores long pedicellate.

Peridium pallid tan, finely tomentose ............... B. verrucosa G. H. Cunn.
Peridium dark brown, areolate ............... B. bovistoides (C. et M.) Lloyd
Sporeseapedicellater a weed epee cout ahs cksvewoss| Ce ee ee ER een: B. pusilla Lioyd

4. Bovista Dill., ex Persoon.
(Descriptions, Proc. LINN. Soc. N.S.W., 1925, 1, p. 369.)

GEyouliiniierr soll Soba ceaaododal.coddnoadaocDddoboooaoco0o dU DOS B. brunnea Berk.
Capillitivumenotepiteede yee eiccsierst. sonc</clchsweleee manuel eae eer HOn CRORE ie B. purpurea Lloyd
5. DISCISEDA Czern.

(Descriptions, Proc. LINN. Soc. N.S.W., 1927, lii, p. 236.)
Spores long pedicellate.

Sporessstroneliy | VeELUCOSC amiarietel here oraieraer ne cerenens D. pedicellata (Morg.) Hollos.
Spores verrucose-areolate .................-- D. hyalothrix (C. et M.) Hollos.
Spores apedicellate (or apiculate, stumps only of pedicels persisting).
Sporesalmostsmoothipsea. see oo iie eee D. candida (Schw.) Lloyd
Spores finely verruculose.
Stoma fimbriate-mammose ............... D. cervina (Berk.) G. H. Cunn.

Stoma tubular, surrounded by a depressed groove.
Spores finely verrucose-echinulate .. D. anomala (C. et M.) G. H. Cunn.
Spores covered with fine round-topped warts .. D. australis G. H. Cunn.
Spores coarsely verrucose.
Spores covered with flat-topped echinulations) {422252626 sso. + s+ be oe ee cele
ERG OLE GEN EGES OPE GB OSOT ONE: CRG IERD- CIS oo Oto D. hypogaea (C. et M.) G. H. Cunn.
Spores covered with coarse verrucae .......... D. verrucosa G H.

6. Apstoma G. H. Cunn.
(Descriptions, Proc. LINN. Soc. N.S.W., 1927, lii, p. 241.)
Gleba purple; spores finely reticulate ........ A. purpureum (Uloyd) G. H.
Gleba olivaceous; spores coarsely reticulate .......... A. reticulatum G. H.
MYCENASTRUM Desv.
(Description, Proc. LINN. Soc. N.S.W., 1927, lii, p. 245) .. M. corium (Guers.) Desv.

8. GEASTER Mich.
(Descriptions, Proc. Linn. Soc. N.S.W., 1926, li, p. 76.)
Mouth peristomate.
Peristome sulcate.
Exoperidium not hygroscopic.
Endoperidium pedicellate.
Endoperidium smooth (or farinose, not roughened).
Peristome circular. 2
Base of the endoperidium smooth or striate ..............
A LP RCC ERT AG oC G. pectinatus (Pers.) Lloyd
Base of the endoperidium plicate ...... G. plicatus Berk.
Base of the endoperidium with a collar-like ring ........
ED) ARE PTE TERERRCIG IG OF CY Ach C1 OCR PRCREE ROS G. Bryantii Berk.

Peristomerellipricaliva tiie ceciiekeets eee G. ellipticis G. H. Cunn.
Endoperidium roughened-verrucose.
Peristome Concolorous! 20s eee ree ranore) sro G. Hariotii Lioyd

Peristome surrounded by a silky, differently coloured zone ....
PS PE RO OCTOBRE EIEN. 9-0 Ci. OO OND U REIL G. Berkeleyi Mass.
Endoperidium sessile or sub-pedicellate (G. Hariotii).

BY G. H. CUNNINGHAM. 255

Exoperidium hygroscopic.
Endoperidium pedicellate.

SORES) Hak Chey, osocconouddcoUDODG OCOD AOOS G. canvpester Morg.
Spores 4-5-5 mw diam.
HMndoperidiumimasperatey craic.) <i cielelelecielene G. Clelandii Lloyd
WINAOPeridivIMey SMO Oth ae aieveseiecle ctecneereeeees G. Smithiu Lloyd
Endoperidium typically sessile ................ G. Drummondii Berk.

Peristome fibrillose.
Exoperidium not hygroscopic.
Endoperidium pedicellate.
Plantsktypica lily sminiuitewe eee e G. minus (Pers.) G. H. Cunn.
Plants ty picalliya lars cuentas rereiehiclcicreke G. limbatus Fries
Endoperidium sessile.
Exoperidium externally felted-tomentose or tomentose-strigose.

Plants lar sews) = Gi Cmbarervocialaenekarerevors eieielisy ce) eve G. velutinus Morg.
Plants smajllae5=2 eerie prercrcsrreeienekclshe heres G. mirabilis Mont.
Exoperidium externally smooth, or almost so.
Spores smooth or nearly so .... G. swbiculosus Cke. et Mass.
Spores verrucose-echinulate or verrucose.
syooey) Aoiasto) f, Chiehi “Goocascouagbaadou G. saccatus Fries
Siders: 4 7 Chievo, congancugonooun0c006 G. triplex Jungh.
Sodas) Yas 7 Chev, soooaqnccsn0noodo00dD G. australis Berk.
Exoperidium hygroscopic.
Hndoperidiumy pedicellaten mmm ceyieecieeteecieacicicleeiens G. arenarius Lloyd
Hindoperidiummesessileman sacs neoo iio aisitioariciene (G. mammosus)

Mouth naked.
Exoperidium not hygroscopic.
Endoperidium pedicellate.

Exoperidium typically fornicate .... G. fenestriatus (Pers.) G. H. Cunn.
HI XODeTICINIMNELeVOlULCMA eI ee eee (G. rufescens)
MING OPEFIGIUM VSESSIC Mas hoes ONGC eae Toler Ae Deon eae aioe G. fimbriatus Fries
Exoperidium hygroscopic. .
SPORES MO 1 ies IAT Ne eee yt RM OM ER Nery. calor An Wen auc are dalaltapal 6 G. floriformis Vitt.
SOO 1-H) (5 Chey, scooochuonassoo0ouncbanoOBDasaUbaNoDG G. simulans Lloyd
Synonyms Recorded in Australasian Literature.
Bovista
anomala Cke. et Mass. = Disciseda anomala
argillacea Pat. = Disciseda cervina
aspera Lev. = Lycoperdon asperum
candida Schw. = Disciseda candida
cervina Berk. = Disciseda cervina
circumscissa Berk. et Curt. = Disciseda candida
debreciensis (Hazsl.) de Toni = Disciseda cervina
gigantea (Pers.) Nees = Calvatia gigantea
hyalothrix Cke. et Mass = Disciseda hyalothrix
hypogaea Cke. et Mass. = Disciseda hypogaea
lilacina Berk. et Mont. = Calvatia caelata
Muelleri Berk. = Disciseda Muelleri
olivacea Cke. et Mass. = Calvatia candida
ovalispora Cke. et Mass. = Bovista brunnea, probably
pusilla Pers. = Lycoperdon pusillum
suberosa Fries = Mycenastrum corium
subterranea Peck = Disciseda cervina
tomentosa Vitt. = Misdetermination
tunicata Bon. = Calvatia candida
Bovistella
aspera (Lev.) Lloyd = Lycoperdon asperum
australiana Lloyd = Lycoperdon glabrescens
cuprica Lloyd = Nomen nudum
glabrescens (Berk.) Lloyd = Lycoperdon glabrescens
Gunnii (Berk.) Lloyd = Lycoperdon Gunnii
nigrica Lloyd = Lycoperdon scabrum
rosea Lloyd = Lycoperdon glabrescens

scabra Lloyd = Lycoperdon scabrum

256 GASTEROMYCETES OF AUSTRALASIA, ix,

Calvatia

aurea Lloyd ; Calvatia candida var. rubro-flava
cyathiformis (Bosc.) Morg. Calvatia lilacina

favosum (Bon.) Lloyd Calvatia caelata

Fontanesii (Dur. et Mont.) Lloyd Calvatia caelata

maxima (Schaeff.) Morg. Calvatia gigantea

occidentalis Lloyd Calvatia candida

olivacea (Cke. et Mass.) Lloyd Calvatia candida

primitiva Lloyd Calvatia gigantea

rubro-flava (Cragin) Lloyd Calvatia candida var. rubro-jlava
Sinclairii (Berk.) Lloyd Calvatia caelata

Oe SW

it]

Cycloderma
ohiense CkKe.
platyspora CkKe. et Mass.

Geaster velutinus
Geaster velutinus

No

Catastoma
anomalum (Cke. et Mass.) Lloyd = Disciseda anomala
circumscissa (B. et C.) Morg. = Disciseda candida
debreciensis (Hazsl.) Hollos = Disciseda cervina
hyalothrix (C. et M.) Lloyd f = Disciseda hyalothria
hypogaea (C. et M.) Lloyd = Disciseda hypogaea
magnum Lloyd = Nomen nudum
Muelleri (Berk.) Lloyd = Disciseda Muelleri
pedicellata Morg. = Disciseda pedicellata
purpurea Lloyd = Abstoma purpureum
subterranea (Peck) Morg. = Disciseda cervina

Disciseda
circumscissa (B. et C.) Hollos = Disciseda candida
debreciensis (Hazsl.) Hollos = Disciseda cervina

Geaster
affinis Col. = Geaster triplex, probably
Archeri Berk. = Geaster triplex
argenteus Cke. = Misdetermination
asper (Mich.) Lloyd = Geaster campester
biplicatus Berk. et Curt. = Geaster plicatus
caespitosus Lloyd = Geaster mirabilis
calceus Lloyd = Geaster minus
coriaceus Col. = Geaster triplex
coronatus Col. = Geaster triplex, probably
coronatus (Schaeff.) Lloyd = Geaster minus
delicatus Morg. = Geaster floriformis
dubius Berk. = Geaster velutinus
Englerianus Henn. = Geaster triplex
fornicatus (Huds.) Fr. = Geaster fenestriatus
fornicatus Fr. = Geaster minus
granulosus Fcl. = Geaster minus
hungaricus Hollos = Geaster floriformis
hygrometricus (Pers.) Fr. = Misdetermination
involutus Mass. = Geaster Drummondii
lageniformis Vitt. = Geaster triplex
lignicola Berk. = Geaster mirabilis
Lloydii Bres. et Pat. = Geaster velutinus

Geaster minus
Geaster minus

marginatus Vitt.
minimus (Schw.) Lloyd
Morganii Lloyd Geaster triplex
orientalis Hazsl. Geaster Bryantii
pseudomammosus Hollos. = Geaster campester
veaderi Cke. et Mass. Geaster velutinus
Schmidelii Vitt. = Geaster pectinatus
striatus Fr. Geaster pectinatus
striatulus Kalch et Cke. Geaster Drummondii
tenuipes Berk. Geaster plicatus

out Il

ii

BY G. H. CUNNINGHAM.

Geaster
tunicatus Vitt.
umbilicatus Quel.
violaceus Lloyd

Lycoperdon
australe Berk.
Bovista Fr.
bovistoides Cke. et Mass.
candidum Pers.
candidum (Rostk.) Bon.
cepaeforme (Bull.) Mass.
Colensoi Cke. et Mass.
coloratum Peck
Cookei Mass.
coprophilum Mass.
corium Guers.
cruciatum Rostk.
dermoxanthum Vitt.
favosum Bon.
Fontanesii Dur. et Mont.
furfuraceum Schaeff.
gemmatum Batsch
giganteum Batsch
hiemale Vitt.
hungaricum Hollos
lilacinum (D. et M.) Mass.
macrogemmatum Lloyd
microspermum Berk.
natalense Cke. et Mass.
nigrum Lloyd
novae-zelandiae Lev.
piriforme var. flavum Lloyd
pseudopusillum Hollos
pratense Pers.
purpuraceum Berk.
purpureum Lloyd
reticulatum Berk.
retis Lloyd
rubro-flavum Cragin
semi-immersum Lloyd
Sinclairii Berk.
substellatum B. et Curt.
tasmanicum Mass.
tephrum Berk.
violascens Cke. et Mass.
Wrightii Berk.

Mycenastrum
bovistoides CkKe. et Mass.
chilense Mont.
clausum Schulz.
leptodermeum Dur.
olivaceum Cke. et Mass.
phaeotrichum Berk.
radicatum Dur.
spinulosum Peck

Scleroderma
bovistoides (Cke. et Mass.) de Toni
chilense (Mont.) de Toni
corium Grev.
leptodermeum (D. et M.) de Toni
olivaceum (Cke. et Mass.) de Toni
phaeotrichum (Berk.) de Toni
radicatum (Dur.) de Toni
spinulosum (Peck) de Toni

i]

257

Geaster fimbriatus
Geaster pectinatus
Geaster triplex

Lycoperdon asperum
Calvatia gigantea
Bovistella bovistoides
Misdetermination
Calvatia candida
Lycoperdon polymorphum
Lycoperdon perlatum
Lycoperdon polymorphum
Lycoperdon spadicewm
Bovistella coprophila
Mycenastrum corium
Lycoperdon candidum
Lycoperdon pusillum
Calvatia caelata

Calvatia caelata

= Lycoperdon polymorphum

Lycoperdon perlatum

= Calvatia gigantea

oul

Misdetermination
Lycoperdon polymorphum

= Calvatia lilacina

Lycoperdon perlatum

= Lycoperdon pusillum

Lycoperdon depressum
Lycoperdon polymorphum
Calvatia lilacina
Lycoperdon piriforme
Lycoperdon pusillum
Unknown
Misdetermination

= Lycoperdon purpuraceum

= Mycenastrum

I

= Mycenastrum

iT]

Hou al

Lycoperdon pusillum, probably
Doubtful species

Calvatia candida var. rubro-flava
Lycoperdon pusillum

Calvatia caelata
Misdetermination

Lycoperdon perlatum

Lycoperdon subincarnatum
Calvatia lilacina
Misdetermination

Bovistella bovistoides
Mycenastrum corium
corium
coriun
corium
corium
corium
corium

Mycenastrum

Mycenastrum
Mycenastrum
Mycenastrum

Bovistella bovistoides
Mycenastrum corium
Mycenastrum corium
Mycenastrum corium
Mycenastrum corium
Mycenastrum corium
Mycenastrum corium
Mycenastrum corium

THE XEROPHYTIC STRUCTURE OF THE LEAF IN THE
AUSTRALIAN PROTHACHAE. Part i.

By A. G. HAMILTON.
(Plates xxiii-xxv and 27 Text-figures.)

[Read 31st August, 1927.]

The endemic Angiosperm flora of Australia is, for the most part, strongly
xerophytic, and this characteristic reaches its highest development in the
Proteaceae, every species, even those growing in the coastal rain-forests, showing
xerophytic adaptations in some degree, the genera Isopogon, Petrophila, Grevillea,
Hakea, Banksia and Dryandra being highly specialized in this direction.

Most of the recognized devices for checking transpiration are found in the
plants of this family—alteration of the orientation of the leaf, reduction of leaf
surface, thickening and cutinization of the epidermis, occurrence of hypoderm,
sunken stomates, palisade cells in two rows and closely packed, reduction of
intercellular spaces, larger water-carrying vessels than in mesophytes, an enormous
development of sclerenchyma and the presence of tannin in considerable quantities.
In some species the leaf margin is recurved so as to form, with the midrib, two
parallel grooves, which are lined with hairs. The stomates are placed in these
grooves, and the hairs not only protect against heat, but check transpiration by
surrounding the stomates with a still atmosphere saturated with water vapour. In
another group of species the stomates are placed in balloon-shaped cavities on the
underside of the leaf; these cavities are lined with hairs, the filiform extremities
of which form a plug to the orifice, thus preventing the free interchange of air.

Reduction of leaf-surface is very common, but it is not carried to the extreme
of actual leaflessness as it is in many other families. Most species have a coating
of two-armed hairs in the juvenile stage of the leaf, but as a rule the arms drop off
as the leaf develops, leaving the base embedded in the thickening epidermis. But
no species has a dense coating of hair persisting throughout the life of the plant as
in the South African Leucadendron. Adenanthos cygnorum is the only Australian
Protead I know which has a permanent coating of hair, and it is much thinner
and woollier than Leucadendron. Some of the Grevilleas have a dense coat of hair
on the underside of the leaf, which is persistent.

Most of the species have several of these adaptations in their leaves. Coatings
of bloom, wax, varnish or hygroscopic salts are absent, so far as I know, in the
Australian Proteaceae, and ethereal oils do not occur—indeed I am doubtful if
the presence of these is a xerophytic adaptation, although many desert plants
possess them. Secretory cavities are found in Franklandia, and I have recently
seen cavities like oil-glands in a species of Conospermum.

The conditions causing xerophytism are many—excessive heat, intense light,
small rainfall, drying winds, great cold, hot water, as near volcanic springs, light

BY A. G. HAMILTON. 259

or poor soils or those containing salt or other substances injurious to plant life (as
vegetable acids derived from humus) are all concerned in bringing about these
adaptations.

Some xerophytes, when grown in moist conditions, respond by producing
foliage of a mesophytic type, probably a reversion to ancestral form, as in the well-
known case of Ulex Europaeus. But the Proteaceae have their characters so firmly
fixed that they do not alter even under the most favourable circumstances.
Alterations do certainly occur, but very rarely. For instance I once found a plant
of Hakea pugioniformis growing on a sand-dune almost at the edge of the sea,
which had the leaves greatly thickened and succulent; but this was merely adding
another xerophytic character to those already possessed by the plant.

Mr. L. Rodway, Government Botanist of Tasmania, in a letter informs me that
the Tasmanian Proteaceae grow in any soil, wet or dry and in any physical-or
chemical condition. “Banksia marginata grows in any soil, wet or dry, and in any
chemical or physical condition. Hakea lissocarpha, H. macrocarpa, and H. epiglottis
are almost marsh plants. H. pugioniformis will struggle along in dry places but
prefers plenty of moisture. Grevillea australis grows in wet places at a high
altitude. In fact, with the exception of Persoonia juniperina, which prefers sandy
heaths, the Tasmanian Proteaceae flourish in wet places.’’ Yet in the material
which Mr. Rodway sent me I can detect no modification or diminution of the
xerophytic characters. Probably the soil conditions and the presence of vegetable
acids account for the retention of these characters, but I am inclined to think
it is due to heredity—the characters have been so long fixed that no alteration in
environment will affect them.

I saw a notable instance in Banksia integrifolia, growing in the rich basaltic
soil of a gully between Mt. Wilson and Mt. Irvine. In the sandy soil near the
coast the tree has cuneate leaves not more than six inches in length, the margin
usually entire, but sometimes slightly serrate. The upper surface is smooth and
dull green. So far as I have seen the trees in this habitat rarely exceed thirty
feet in height. But at Mt. Wilson they are at least seventy feet high. In full
grown trees the leaves are similar to those of the coast tree, but in young trees
from ten to twelve feet in height they are lanceolate, eighteen inches long, very
dark green and shining on the upper side, and have a strongly serrated margin.
This is the most remarkable instance of alteration brought by environmental change
that I have seen but it was an alteration in external form only—sections of the
leaves showed the characteristic structure of the coast plant.

In Hakea trifurcata and several other species with terete leaves, the plants
bear reversionary leaves in their ordinary habitat. The bulk of the leaves are
terete, but the plants have large numbers of flat leaves up to three inches in
length, and elliptical to spathulate in shape. This is not due to any change of
environment, for in their native habitat the flat leaves are just aS numerous as
they are under cultivation.

Professor J. B. Cleland some years ago showed me a very peculiar example
of alteration in a plant of Hakea vittata which he had grown from seed in his
garden. The plant at first had terete leaves, but after a time flat leaves appeared,
which seems to be the opposite of what one would expect. Hakea orthorrhyncha
also shows a considerable amount of variation, as will be seen later. But the
altered leaves in every case are just as xerophytic in structure as the ordinary
leaves and show no approach to the structure of a mesophytic leaf.

H

260 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, i,

Notwithstanding their very perfect adaptation it is very noteworthy that the
Proteaceae are very sparsely distributed in the arid and semi-arid regions of
Australia, both in regard to species and individuals. For instance, in the Broken
Hill district only three of the family have been found, and these in small numbers,
while the apparently less efficiently protected Hucalypts, Acacias, Hremophilas
and Salt-bushes are numerous both as to species and individuals. The same
paucity of Proteads is noticeable all over Central Australia. It is also remarkable
that these members of the family which are most highly specialized (such as
Banksia and Dryandra) do not occur in these dry regions. The stronghold of
the family is the belt of well-watered country bordering the Continent, and
possessing a sandy soil. It therefore seems to be the edaphic factor which governs
the distribution of the Proteaceae—they require a highly siliceous soil.*

Mr. E. Cheel has recorded® the presence of what may be possibly mycorrhizae
on the roots of some of these plants, and if this is correct it is probable that it
accounts for their scarcity in the dry interior—the fungi may not be able to grow
under these extreme conditions.

Genus HAKEA.

For the purpose of this paper, the Hakeas may be divided into two groups—
those with flat and those with terete leaves. Both groups contain species with
entire and with divided leaves. As before noted in some species it is common to
find both on the same plant. There are a few species which fall between the two
main groups; they are flattened but very narrow. These I shall deal with among
the terete species.

The flat-leaved species have a vertical orientation, and are all centric. A few,
as H. cucullata and H. auriculata, are not vertical, but the structure is also centric.
The leaves are thick (but in varying degree), flat and leathery. The margin may be
entire or toothed, and in the latter case the tooth ends in a prickle formed by the
vein running through it. A few, as H. ceratophylla, have divided leaves. The
surface of the leaf is generally dull, but there is no coating of bloom, wax or
varnish, even on the young leaves. There are only a few species with a persistent
coating of hairs, and that is very sparse, but all are hairy in the juvenile stage.
The hairs are T-shaped (Text-fig. 1), the pedicel being very short. They are
thick-walled, with a very narrow lumen, and are silvery or ferruginous in colour.
The arms drop off early in the life of the plant, but the pedicel persists and
later becomes embedded in the epidermis. In a paper on H. dactyloides® I
spoke of these as capitate hairs, but further investigation showed their real
nature. Solereder™ says that these hairs are probably found in every species of
Hakea and Grevillea, and my experience confirms that opinion, as I have found no
species of either of these two genera without them. The epidermis is thick, with
highly cutinized cuticle, the thickness increasing with age. The thickness of the
epidermis varies from 0:02 mm. in H. undulata to 0-103 mm. in HF. incrassata.
The cuticle is from 0-003 in H. undulata to 0:04 mm. in H. ceratophylla. The
a ee Ee ee ee aes

* Since this was written, I have read the recent paper by McLuckie and Petrie, “An
Ecological Study of the Flora of Mount Wilson, Pt. iv’ (These PROCEEDINGS, 1927,
pp. 161-184) in which the authors state that the main contributing factor in xerophytism
is the low water-retaining capacity of light sandy soil, and on consideration, I think
that this is probably the case. It must be remembered, however, that in no locality, so
far as my experience goes, are Proteads plentiful on non-sandy soil, wet or dry, while

on the other hand, they are found in numbers on sandy soils which are always wet and
even swampy.

BY A. G. HAMILTON. 261

lateral walls of the cells are thin and the cavities are usually empty, but sometimes
contain crystals of calcium oxalate, tannin, or in a few cases, a substance staining
faintly with picric-anilin blue. The outer surface of the cuticle is often ridged
and there is sometimes a ridge-like thickening in the lateral walls of the cells
surrounding the orifice of the air-chamber above the guard cells (Plate xxiii,
fig. 1). The cells which lie round this cavity (which I shall call the vestibule)
in many species project above the general surface of the leaf; there is thus formed
a crater-like opening to the stomata. Hypoderm, in the ordinary sense of the word,
does not occur, but in a few species a sheet of sclerenchymatous fibres lies just
under the epidermis.. The vestibule may be bell, dome, funnel, or hourglass
shaped (Text-figs. 2-8). In H. corymbosa it is hourglass shaped, and Solereder“
says that in H. cyclocarpa it is double-funnel shaped (Text-fig. 5), but in my
sections it is cylindrical, somewhat constricted in the middle, but not so much as
to be hourglass shaped. ‘In H. cucullata I observed a few much constricted in the
middle (Text-fig. 7), but the majority were the usual type. The guard-cells are
at the bottom of the vestibule on a level with the top of the palisade cells.

ee
TE TOM
DC D DO

: on Beans

Text-figures 1-8. 1. Two-armed hair. 2-8. Vestibules of stomata.
2, H. macrocarpa; 3, H. Hookeriana; 4, H. arborescens; 5, H.
amplexicaulis; 6, H. megalosperma; 7, H. cucullata; 8, H. linearis and
H. ruscifolia. The figures are diagrammatic.

The vascular bundles run through the central region; they are often prominent
on the lower or on both sides of the leaf. They vary considerably in number, size,
position in the mid-region, and in the amount of sclerenchyma fibres which
accompany them. In most species the conducting tissue is in very small proportion
to the sclerenchyma fibres which enclose it.

The most characteristic feature of the leaves of the genus is the enormous
development of sclerenchyma. It occurs in three forms—fibrous, columnar, and
massive. Fibrous sclerenchyma is found in every member of the genus. It usually
occurs in connection with the vascular bundles, a strand running along the upper
and lower sides of the bundles, and in some cases all round. Sometimes a sheet

262 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, i,

of fibres lies just under the palisade, a feature in which they resemble the
Banksias. The large amount of sclerenchyma has a very marked stiffening effect
on the leaf.

The columnar variety is almost general in the genus, a few exceptions
occurring among both flat and terete leaved species, but in some of the flat-leaved
forms there are only a very few columns. The columns are cylindrical, spreading
out at the top and bottom like the base and pediment of a pillar (Text-fig. 17).
In some species the extensions are so wide that they form an almost continucus
sheet between the palisade and the epidermis (Text-fig. 19), and sometimes
spikes or pegs grow up from this through the lateral walls or even through the
cavities of the cells (Text-fig. 19). At the lower end they often send off root-like
processes, which penetrate the mid-region and sometimes unite with the outer row
of storage cells, thus thickening their walls very considerably. Some of the
massive sclereides (when they occur among the palisade cells) appear to be derived
from the columns of secondary growth but those in the mid-region are of indepen-
dent origin. They are of irregular and extraordinary shapes (Text-figs. 9-16). The
amount of sclerenchyma in the leaves of the Hakeas is very large—some species as
much as 50%, and in H. multilineata and H. obtusa even more.

wy ee
F spel

Text-figures 9-16. Forms of detached sclereides. 9-13 and 16,
H. corymbosa; 14 and 15, H. cucullata.

There are several distinct types of leaf structure although there is a general
resemblance all through the flat-leaved Hakeas. Solereder’’ quotes J6Onsson’s
classification of the leaf structure of the Proteaceae from which the following is
an extract.

I. Hakea-type. Leaf structure centric; sclerenchyma e¢ells in palisade tissue.
Species of Adenanthos, Grevillea, Hakea, Isopogon, Molloya, Petrophila,
Roupala, and Stenocarpus.

Il. Isopogon-type. Leaf structure centric; ordinary sclerenchyma cells. Species
of Bellendena, Hakea, Isopogon, Leucadendron, Nivenia, Sorocephala and
Xylonelum.

V. Synaphea-type. Leaf centric; sclerenchyma of the vascular bundles vertically
transcurrent. Species of Hakea and Synaphea.

BY A. G. HAMILTON. 263

There are, however, many other differences among the various species of
flat-leaved Hakea, and I propose to arrange them according to these peculiarities, as
follows:

Group I. Species with thick leaves; palisade in two rows; eentral region with
thick-walled cells; columnar and fibrous sclerenchyma both present (Plate
Soa, 1S, Uo =

H. macrocarpa A. Cunn. The leaf in this species is thick and smooth.
Bentham™ says ‘“Minutely pubescent on both sides’, but I have not been able to
detect pubescence in the mature leaves in herbarium specimens’ In sections,
however, the persistent bases of the hairs are seen embedded in the epidermis.
The leaf is 0-52 mm. thick, in the thinner places between the veins, and 0-84 mm.
at the prominent midrib.

The epidermis is 0:05 mm. and the cuticle 0:03 mm. The lateral walls of
the cells are thin and they often contain tannin. There are stomates on both
sides, and the epidermal cells surrounding the stomatal opening are slightly raised
above the general surface of the leaf. The vestibule is dome shaped (Text-fig. 2)
and is 0:06 mm. wide, and the guard cells are at the level of the top of the
palisade cells. There are two rows of palisade cells with a total depth of 0-16 mm.,
and they are closely packed except where they open out in an air-chamber under
the guard-cells. The lower row is continuous under the air-chamber. The mid-
region is from 0-24 to 0:28 mm. across and is made up of thick-walled cells
frequently containing starch and sometimes tannin. Columnar stereides run
through the palisade; they are slender and rather sparsely distributed; they
spread slightly under the epidermis and send out root-like processes which coalesce
with the walls of the outer layer of storage cells beneath, thickening them very
much (Text-fig. 17).

Five large veins run through the mid-region, accompanied on both top and
bottom by narrow strands of sclerenchyma fibres, the diameter of the whole
bundle being 0-4 mm., while the conducting part of the vein is 0-3 mm. On the
ventral side the cord of fibres is not so thick as on the dorsal. Besides these five
veins there are many smaller ones just beneath the palisade on the ventral side;
these have no accompanying strand of sclerenchyma.

H. arborescens R. Br. This has rather a thick leaf, ranging from 0-56 to
0.7 mm. The epidermis is ridged; on the greater part of the leaf it is 0:04 mm.
thick, but on the margin it thickens up to 0:07 mm. The cuticle is 0-016 to 0:04 mm.
thick, and is highly cutinized. The vestibule is conical (Text-fig. 4) and does
not project above the general surface of the leaf; it is 0:03 mm. wide. The palisade
is in two rows and has a total depth of 0:16 mm. There is a great development of
sclerenchyma. The columnar stereides are 0:02 mm. thick, and spread out
slightly at the top; they send out root-like processes at the base but these do not
unite with the walls of the storage cells as they do in some other species
(Text-fig. 22 and Plate xxiii, fig. 2). The central region is 0:24 to 0:26 mm. wide,
and consists of thick-walled cells, many filled with tannin in which starch grains
are embedded. The veins are numerous but small, the largest being 0:3 mm. in
diameter and the conducting tissue 0:16 mm., but the majority are much smaller
than this.

H. cyclocarpa Lindl. Leaf thick, 0:64 mm., epidermis 0:04, and cuticle
0:02 mm. The palisade is in two rows, closely packed and 0:14 mm: deep. The
central region is made up of thick-walled cells and is 0:14 mm. across. The veins
are small and sub-equal, 0-2 mm. in diameter and the conducting tissue 0-12 mm.
There are many columnar sclereides spreading out beneath the epidermis and

264 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, ils

above the central region. Throughout the central region there are many bundles
of sclerenchyma fibres apart from those capping the veins.

H. Roeii Benth. Leaf thick, 0:74 mm. Epidermis 0:06 mm., and the cuticle
0:04 mm.; vestibule 0-18 x 0:14 mm., with the edges of the orifice projecting above
the surface of the leaf. The palisade is two rows 0-16 mm. deep and the cells
0:018 mm. in diameter. The columnar stereides are very numerous, and spread
a little at the top and bottom, the outgrowths uniting with walls of the storage
cells of the middle region (Text-fig. 22). This last is 0-42 mm. deep and is
composed of thick-walled cells, many filled with starch and a few with tannin
scattered irregularly about. The vascular bundles are numerous but small, the
midrib being very little larger than the others; it is 0-32 mm. in diameter and the
conducting tissue 0:14 mm. They all have caps of sclerenchyma on both ends and
these extend laterally so that the bundles are almost entirely enclosed. Just
beneath the palisade (but separated from it by a single layer of thick-walled cells)
there are many bundles of sclerenchyma fibres.

H. Hookeriana Meiss. Leaf thick, 0-72 mm., the epidermis 0:04 mm., and the
cuticle 0:02 mm. The vestibule is bell-shaped (Text-fig. 3) and 0-06 x 0:04 mm.
There are two rows of palisade cells with a total depth of 0-24 mm. The columns
are numerous and spreading at top and bottom. At the base they form irregular
masses, penetrating the mesophyll. This last consists of thick-walled cells, but
not so thick as in the preceding species; it is 0-3 mm. across. The veins are few
and very small, the largest being only 0:06 mm. in diameter and the conducting
tissue 0:04 mm.

H. incrassata R. Br. The leaf is 0:74 mm. thick, with a smooth surface.
Epidermis 0-05 mm. and cuticle 0:01 mm. thick. Palisade in two rows, 0:14 mm.
deep. The medullary tissue is 0-28 mm. across and made up of thick-walled cells,
very many filled with starch or tannin. The columnar stereides are very
numerous and closely spaced; they often join at the base. Bundles of fibres lie
almost continuously under the palisade, but separated from it by a layer of storage
cells. The veins are few, their long diameter is 0-36 mm. and the conducting
bundle 0-08 mm.

H. flabellifolia Meiss. The leaf of this species resembles that of H. Brownii
closely both in external form and in internal structure. The veins are, perhaps,
a little less in diameter, and the epidermis a little thicker. The leaf is 1-6 mm.
thick; the epidermis 0-08 mm. and the cuticle 0:02 mm. The palisade is in two
rows, the cells rather slender with a depth of 0:18 mm. The central region is
composed of thick-walled cells, many filled with tannin, it is 0-32 mm. across.
The veins are rather small, 0-28 mm. in diameter and the conducting tissue 0-14
mm. There are numerous columns but perhaps not so many as in H. Brownii, and
they do not spread out so much at the top and bottom as in that species.

H. Brownii Meiss. The leaf is very thick, 0:92 to 1 mm., the epidermis
0:06 and the cuticle 0:02 mm. ‘The vestibule is conical and rather wide (Text-
fig. 4). The palisade is in two rows and closely packed. It is 0-2 mm. deep.
Thick-walled storage cells occupy the midregion which is 0-4 mm. across; many
of these cells are filled with tannin. The veins are 0°5 mm. in diameter and the
conducting tissue is 0-2 mm. A very large number of stereides traverse the
palisade so that at least 50% of that region is composed of sclerenchyma. The
tops and bases of these spread out a good deal so that there is a continuous sheet
under the epidermis in places. Here and there small masses of sclerenchyma
develop in the cavities of the epidermis (Plate xxiii, fig. 4).

BY A. G. HAMILTON. 265

H. Bacteri R. Br. The leaf is 0:74 mm. thick, with a smooth surface but the
cells round the orifice of the vestibule project above the surface. The epidermis
is 0-07 mm. and the cuticle 0-025 mm. thick The vestibule is bell-shaped and is
0:06 x 0-08 mm. The air-chamber beneath the guard cells goes the full depth of
the two palisade layers, which have a total depth of 0:12 mm. The middle of the
leaf is occupied by thick-walled cells; it is 0°32 mm. wide; many of the cells are

19
QO000 OoOQoone

TEES sh.

A — va AR — co
ER

Text-figure 17. Extension of base of column thickening walls of
storage cells. 18. Section perpendicular to axis of vein, bases of
columns enclosing vascular bundle. 19. Spike-like extensions from
sclerenchyma penetrating the epidermis; H. ruscifolia. 20. H. ambigua,
columns sending root-like extensions between the two rows of palisade
cells. 21. Section parallel to plane of leaf, H. amplexicaulis, column
extensions enclosing vascular bundle. The figures are diagrammatic.

ep. = epidermis, vb. = vascular bundle, col. = columnar stereides,
sh. = sheet of sclerenchyma formed by coalescence of tops or bottoms
of columnar sclereides. :

full of starch or tannin or both. A few thick veins run through this part, 0:4 mm.
in long diameter, the conducting tissue being 0:24 mm. The columnar stereides are
not numerous; they spread out at the top, forming a sheet under the epidermis;
at the base they send down extensions into the mid-region and these frequently
coalesce with the storage cells of the outer layer, thickening them very much.

H. ceratophylla R.Br. This species is notable for the extreme range of shape
in the leaf, which is divided. It is 0-9 mm. thick, with a slightly ridged surface.
The epidermis is 0:06 mm. and the cuticle 0:04 mm. in thickness. The vestibule is
conical and slightly depressed below the leaf surface; it is 0:02 mm. x 0:04 mm.
The palisade is in two rows, the cells rather slender, and it has a total depth of
0-16 mm. The medullary tissue is 0:5 to 0:54 mm. thick, and consists of thick-

256 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, i,

walled cells which are filled with starch, those surrounding the veins are full
of tannin (Plate xxiii, fig. 3). The stereides are thick—0-:04 mm. They expand
at the top, forming a sheet under the epidermis and the lower extremities unite
with the upper layer of storage cells. The vascular bundles extend from the
upper to the lower epidermis, and are heavily capped with sclerenchyma fibres
on both sides.

H. Candolleana Meiss. This species has narrow leaves varying from 1 to 4
mm. in width, but the structure is identical in each. The leaf is 0-44 mm. thick,
epidermis 0:07 mm. and the cuticle 0:025 mm. The vestibule is narrow and conical.

Text-fig. 22. H. amplexicaulis, root-like extensions of columnar
stereides. 23. H. pandanicarpa, section parallel to plane of leaf showing
arrangement of palisade, and sclerenchyma round the stomates. 24.
H. pycnoneura, sectional outline of leaf showing arrangement of veins
and bundles of sclerenchyma fibres. 25. H. Morrisonianwm, section of
leaf, showing arrangement of veins and bundles of sclerenchyma fibres.
26 and 27. H. ambigua, branching columns. (All diagrammatic.)

col. = columnar stereides, ep. = epidermis, ex. = root-like extensions,
fse. = fibrous sclerenchyma, pal. = palisade, sc. = sclerenchyma fibres,
st. = stomate, vb. = vascular bundle.

There are two rows of palisade, 0:14 mm. deep. The central region is made up
of thick-walled cells containing much starch, and very many tannin filled, it is
0-6 mm. across. The mid-vein is 0:24 mm. in diameter, and the conducting tissue
0:14 mm. Unlike most of the other species the sclerenchyma caps do not encroach
on the palisade. The minor and marginal veins are just beneath the palisade, but
the mid-rib is midway between the upper and lower layers. The marginal vein
has a thick strand of sclerenchyma fibres below, and a little to one side. There
are many slender columns which have extended bases but do not spread at all
at the top. More than twenty bundles of sclerenchyma are found beneath the
palisade tissue.

BY A. G. HAMILTON. 267

H. falcata R.Br. The leaf is from 0-6 to 0-65 mm. thick with a ridged
epidermis, the latter 0:08 mm., and a rather thin cuticle, 0:02 mm. The vestibule
is dome-shaped 0:05 mm. wide; the surrounding epidermal walls do not project
above the leaf surface. The palisade is very closely packed and is 0:14 mm. deep.
The central region is 0:2 to 0:25 mm. across; some of the cells are pitted. The
middle vein is 0-5 mm. and the conducting tissue 0°3 mm. There are not many
columns.

H. pycnoneura Meiss. This species and the next resemble H. florida and
H. falcata in the hardness of their leaves but they are also remarkable for their
sectional outline (Text-figs. 24 and 25). Their leaves resemble those of some of
the Grevilleas, and so far as my experience goes, they stand alone amongst the
Hakeas. The leaf of this species is 0-66 mm. thick, and carries a great many two-
armed hairs, the arms of which drop off while the bases persist and become
embedded in the epidermis. The epidermis is very deeply ridged and the cavities
of the cells large; it is 0:06 mm. thick and the cuticle 0.02 mm. The palisade cells
are very tightly packed and only 0:1 mm. deep. The central region is 0-32 mm.
wide, the cells are very thick-walled, a few contain tannin. There is a large
central vein with a heavy capping of sclerenchyma on the dorsal side. This and
the next species differ from the majority of Hakeas in having no columnar
sclereides.

H. Morrisonianum. The central flat part of the leaf is 0-4 mm. thick, while
the dorsal and ventral lobes on each side are 0-48 mm. and 0-44 mm. respectively.
The epidermis is 0:08 mm. thick, thinner on the lobes, and thickly covered with
T-shaped hairs in the juvenile stage. The vestibule is narrow-conical, 0:02 mm.
wide and projects above the surface. The central region is 0-16 to 0-24 mm. wide,
and made up of very thick-walled cells. The palisade is not deep—0-:1 mm.—and
is very closely packed. The veins extend from the floor of the upper epidermis
to that of the lower, and are very heavily capped with sclerenchyma fibres, the
conducting tissue being barely 0-1 mm. As in the last species, there is a central
vein and one in each lobe.

H. florida R.Br. Leaf 0:34 mm., epidermis 0-04 mm., cuticle smooth, 0-016 mm.,
vestibule not projecting, broadly bell-shaped, 0:04 mm. wide. Palisade in single row
0-1 mm. deep. Cells of central region not as thick as in most other species, 0-1 mm.
in diameter. Middle vein 0-5 mm. across and the conducting tissue 0-3 mm.; the
marginal and smaller veins being lightly capped with sclerenchyma.

H. pandanicarpa R.Br. This is a very remarkable species, having an arrange-
ment of the palisade and stereides that I have found in only one other species.
The leaf is thick—0:92 to 0:96 mm.; the epidermis 0:03 mm. and the cuticle 0:02 mm.
The orifice of the narrow bell-shaped vestibule does not project above the leaf
surface. The palisade is in two rows, 0:24 mm. deep. The medullary tissue is
0:32 mm. across and consists of thick-walled cells, many containing starch, and a
few near the middle filled with tannin (Plate xxv, fig. 3). There are a great
many stereides, rather slender, 0:02 mm., spreading a little at the top, and much
more at the base, with root-like extensions which often penetrate the mid-region.
The storage cells of the outer layer have their walls much thickened by union with
these. The veins are few and small, they are capped with bundles of sclerenchyma
on both sides.

The remarkable feature of the leaf is the arrangement of the sclereides and
palisade in relation to the stomates. Under each stomate there is a cylindrical
cavity or air-chamber, surrounded by one row of palisade cells; outside this there

I

268 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, i,

is a circle of columnar stereides two or three rows deep, the tops and bottoms
extending so as to touch each other, while the slender shafts leave openings for
the free interchange of gases (Text-fig. 23). This is the most elaborate provision
against excessive transpiration that I have seen in the Hakeas (Text-fig. 23 and
Plate xxiv, fig. 2). It occurs only in this species and in H. clavata.

H. clavata, Labill. This species does not fit into any of the groups in which
I have arranged the Hakeas, but I have placed it here on account of the arrange-
ment of the stomates, palisade, ete., this being similar to that of H. pandanicarpa
(Plate xxiv, fig. 2). There is, in the herbarium of the Sydney Botanic Gardens,
an undetermined species collected by the late Mr. J. H. Maiden at Esperance, W.A.,
which has the same structure, but as it has no point of difference except a much
shorter leaf, I have no doubt but that it is the same species.

The plant has very thick leaves closely resembling those of Petrophila linearis;
they are 3:2 mm. thick, epidermis 0:04 mm. and cuticle 0:02 mm., vestibule bell-
shaped, 0:06 mm. wide, palisade in two rows 0:32 mm. wide. The central region
is 2-48 mm. across and is composed of very thin-walled cells (Plate xxiv, fig. 3). It
is difficult to say that there is any chlorophyll in them; only herbarium material
was available. At the base of the palisade cells some of the central cells stained a
faint blue which might indicate chlorophyll. These thin-walled cells are much
larger than the thick-walled cells of this group, ranging from 0-06 to 0-15 mm. in
diameter. Bentham describes the leaf as veinless, but there are a few veins, very
small and narrow, and with very little xylem. They are distributed just under
the palisade, and one is in the centre. There is no sclerenchyma anywhere in
the mid-region.

H. obtusa Meiss. Bentham™® says: “The young shoots silky-tomentose, the
adult foliage glabrous’. But in the specimens seen by me, the old leaves were
slightly hairy. 'This species might almost be placed in a section by itself, as the
mesophyll is composed of very thick-walled cells unlike those of any other species
(Plate xxv, fig. 2). This makes it extremely hard, and very difficult to section
without breaking up.

The leaf is 0-6 mm. thick, the epidermis 0:06 mm., and the cuticle rather thick
—0:04 mm. The palisade is in two rows, 0:14 mm. deep. The thick-walled cells
of the mesophyll are rather longer than broad, the long axis being perpendicular
to the plane of the leaf; the small cavities being sometimes filled with tannin and
sometimes with a substance staining faintly with picric-anilin blue; this region
is 0-28 mm. across. The main vein is 0-6 mm. in diameter and the conducting
tissue 0:06 mm. The marginal vein is heavily capped with sclerenchyma. There
are very few columnar sclereides, and those there are seem to arise from the

sclerenchyma of the central region and grow outwards towards the epidermis
through the palisade layer.

Group II. Leaves rather thinner than in the previous group; palisade in one
row (except in H. marginata); cells of mid-region elongated or rounded, and
containing chlorophyll (Plate xxv, fig. 4.)

H. megalosperma Meiss. The leaf is rather thick, 0-5 mm., the epidermis
0-05 mm., ridged, cuticle 0:02 mm. The vestibule opens widely and is constricted
in the middle (Text-fig. 6). The palisade is in two rows and is 0-12 mm. deep.
The veins are small and sub-equal, 0:22 mm. in diameter, with conducting tissue
0-12 mm. There are many columns and they spread widely at the base and apex;
where they are over a vein they almost enclose it.

BY A. G. HAMILTON. 269

H. amplexicaulis R. Br. Leaf 0-4 mm. thick; epidermis 0:04 mm., smooth,
the lateral walls ridged like those shown in Plate xxiii, fig. 1; cuticle 0:02 mm.;
vestibule a narrow reversed funnel, and depressed (Text-fig. 5). Palisade in one row
0-1 mm. deep; central region 0:12 mm. across, made up of thin-walled cells
containing chlorophyll. There is a midrib 0-68 mm. in diameter, the conducting
tissue being 0°5 mm., and a number of smaller veins. The columns are numerous
and spreading at the top and much more at the bottom. Over the veins the
extensions come into contact with each other and form a sheath to the vein
(Text-figs. 18 and 21).

H. glabella R. Br. The leaf is 0-4 to 0-5 mm. thick, the epidermis is 0:04 mm.,
and the cuticle 0-02 mm.; the vestibule is shallow and cylindrical, and the
surrounding cells project above the leaf surface. The palisade is in one row and
is 0-1 mm. deep; the central region is 0:12 mm. across. The columns of
sclerenchyma are few and spread at the top but not at the base. There is one main
vein, 0-56 mm., lightly capped with fibres, and a number of smaller ones, each of
which has a bundle sheath, a feature not common in Hakeas.

H. cristata R. Br. The leaf is 0-52 mm., slightly ridged, epidermis 0:06 mm.,
and cuticle 0:02 mm. The palisade is in one row 0:16 mm. deep; the central region
0-1 mm. across. The stereides spread at both ends and form a partial sheath to
the vascular bundles.

H. linearis R. Br. The leaf is rather thin, 0-38 mm., epidermis 0:06 mm., and
cuticle 0-02 mm.; the vestibule projects so as to form a cone round the orifice, as
also in H. ruscifolia (Text-fig. 8). The palisade is in one row and 0-1 mm. deep;
the central region is 0:12 mm. across. The columnar stereides spread out at the
upper and lower extremities and form a sheath to the vascular bundles as in the
last two species. The main vein is 0-32 mm. in diameter and the conducting vessels
take up 0:24 mm. of this. The marginal veins are small and have a cap of fibres
on the ventral, but none on the dorsal side; a good many tannin cells are near
the veins.

H. ferruginea Sweet. Leaf thin, 0-26 mm., epidermis 0:04 mm., and cuticle
0:01 mm.; vestibule conical, 0:04 mm. wide; the palisade is in one row 0:08 mm.
deep. The central region 0-1 mm. across. A rather unusual feature is the complete
absence of columnar stereides, and there is very little fibrous sclerenchyma.

H. marginata R. Br. The leaf is of medium thickness, 0-42 mm., the epidermis
0:04, and the cuticle 0:01 mm. The vestibule is bell-shaped and does not project
above the leaf surface. The palisade cells are in two rows with a total depth
of 0:08 mm., and are very closely packed. The central region is 0:2 mm., and
consists of thin-walled chlorenchymatous cells closely packed. The main vein is
0-4 mm. in diameter and is completely surrounded by sclerenchymatous fibres, the
conducting vessels taking up 0-2 mm. of this. There is a strong marginal vein,
also surrounded by sclerenchyma. The other veins are small and heavily capped
with fibres on both faces. There are many columns in the palisade, and some
irregular massive ones in the central region, some of the arms of which penetrate
the palisade and take the form of columns.

Group III. Leaves thick or rather thin; palisade in two rows; medullary region
occupied by chlorenchymatous cells arranged in an open network like the
spongy tissue in mesophytic leaves (Plate xxiv, fig. 4).

H. auriculata Meiss. The leaf is 0:36 mm. thick (at the midrib it is 0-86 mm.),
epidermis 0:04 mm., and cuticle 0:02 mm. Vestibule bell-shaped and depressed,
0:04 mm. wide. Palisade in two rows 0-16 mm. deep; central region 0-1 mm.

270 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAB, i,

across. Midrib 0-68 mm. in diameter, the conducting tissue being 0°34 mm. The
small marginal vein has no capping.

H. eriantha R. Br. The leaf is 0-4 mm. thick, epidermis 0-04 mm., cuticle
0:02 mm. Palisade in two rows 0:18 mm. deep. Central region 0-08 mm. across.
The main vein is 0-38 mm. in diameter and the conducting tissue 0:16 mm. The
columns are numerous and spread at the top and bottom; near the veins the
bases enclose them.

The outstanding feature is the narrowness of the mid-region; the palisade and
columns of each side almost meet at their bases.

H. lasiantha R. Br. The leaf is 0:44 mm. thick, epidermis 0:03 mm. The cuticle
is very thin, 0:005 mm. The palisade is two-rowed, 0-1 mm. deep. The spongy
chlorenchyma is 0-1 mm. across. The main vein is 0:54 mm. across, with a
conducting tissue 0-3 mm. There are numerous columns, irregular in shape, and
spreading both at top and bottom; sometimes the inner extensions entirely enclose
a vein, especially the marginal vein.

HA. ruscifolia Labill. Leaf 0-4 mm. thick, epidermis 0:04 mm., and cuticle
0-02 mm.; the walls of the vestibule are raised so as to form a conical projection
above the surface of the leaf, rather higher on the ventral side than on the dorsal,
the cavity being 0:02 mm. wide (Text-fig. 8). Palisade in one row, 0-1 mm. on the
ventral side and 0:06 mm. on the dorsal. Medullary tissue a very open network
0-04 mm. wide. The central vein 0-16 mm. in diameter and the conducting vessels
0-08 mm. The smaller veins are narrowly capped as is the marginal vein.

The columnar sclereides are numerous, many of them curved; they all spread
out at the apex and base. At the apex they form a continuous sheet under the
epidermis, and from this sheet pegs or spikes grow up through the lateral walls
of the epidermal cells and sometimes through the cavities (Text-fig. 19). At the
base the columns also spread out, and where they are over a vein they coalesce
to form a-sheath enclosing it completely (Text-figs. 18 and 21).

H. saligna R. Br. Leaf 0-48 mm. thick, epidermis 0:03 mm., cuticle 0:01 mm.;
vestibule conical, 0:02 mm. wide, and projecting very slightly above the surface
of the leaf; palisade in one row 0-1 mm. deep; central region 0:2 mm. across, and
made up of a rather closely packed network of cells, many, especially near the
veins, being filled with tannin. Veins sub-equal, 0:36 mm. in diameter, the
conducting tissue being 0:16 mm. De Bary, quoted by Solereder, mentions “the
occurrence of solitary crystals in the epidermis”, but I have never seen any in the
sections I have examined, but I find that the occurrence of crystals is often
sporadic, one leaf showing numbers while another from the same plant has none.

H. multilineata Meiss. Leaf 0°38 to 0-4 mm., epidermis 0:08, and cuticle
0:04 mm. thick; vestibule bell-shaped, 0:03 mm. wide; palisade in one row,
yarying in depth 0:2 mm. at most. Medullary region 0:16 mm. wide, the cells small
and very full of chlorophyll. The veins are sub-equal; the conducting tissue is
only 0:2 mm. wide, while the bundles of sclerenchyma run out to the epidermis on
each side with a total depth of 0:74 mm. The columns of sclerenchyma are
numerous and very massive and irregular. Cannon® says, “The sclerenchyma
appears to be mainly or wholly fibrous’, but in all the sections I have examined
there are many columnar stereides. At the margin there is a fan-shaped strand
of fibres enclosing a small vascular bundle. The characteristic feature of the leaf
is the great quantity of mechanical tissue, at least 50% of the leaf substance being
sclerenchyma. This makes it extremely difficult to cut sections. Probably the
habitat of the plant—the arid regions—accounts for this.

BY A. G. HAMILTON. 271

H. laurina R. Br. This species, though growing in the semi-arid parts of
Western Australia, is more mesophytic than any other I have seen, the greater
part of the leaf being chlorenchyma. It is 0°36 mm. thick, the epidermis 0:04 mm.,
and the cuticle 0:01 mm. Vestibule, wide bell-shaped, 0:02 mm. wide. The palisade
is in one row, 0:1 mm. deep; mid-region 0-1 mm. across, and made up of a rather
close network of cells. One central vein 0-44 mm. across and the conducting
tissue 0:12 mm.; the fibres of sclerenchyma almost surround the vein but between
the edges there are a few thick-walled cells containing starch; the other veins are
all small and have very little sclerenchyma, but the marginal bundle is entirely
surrounded by sclerenchyma, the vessel bundle being 0:06 mm. in diameter.
There are no columnar sclereides.

H. eucalyptoides Meiss. This variety has rather thinner leaves than H. lawrina,
and the midrib projects on both sides of the leaf considerably more. But in
other respects it is quite similar.

According to Haberlandt,“” Hintz observed water-storage tissue in the leaf
margin of various Acacia, Quercus, Ilex, and Hakea eucalyptoides. The opinion
is expressed that “The margin, which is the region most exposed to damage by
excessive loss of water, is provided with a local water tissue composed of a
varying number of layers, although water-storage arrangements are altogether
absent, or, at any rate, conspicuously less, in the rest of the leaf’.

The cells referred to are cells between the sclerenchyma cap of the marginal
vein and the epidermis, and they are found at every vein in most species. They
are also found under the epidermis wherever the columns spread out to form a
sheet at the apex. Haberlandt, however, elsewhere, refers to thick-walled cells
as starch-storing tissue. As I have already said, these thick-walled cells may
contain starch, tannin, and crystals, and it is quite probable that they may also
function as water-storing tissue. But they differ much from the typical
water-storing thin-walled cells.

H. corymbosa R. Br. The leaf is 0:32 mm. thick, epidermis 0:04 mm., and
cuticle 0:02 mm.; the vestibule is dome-shaped and 0:06 mm. wide. The palisade
is in one row, 0:08 mm. deep on the ventral side and 0:06 mm. on the dorsal.
Medullary region a very close network, 0:1 mm. across. The middle vein is
0-5 mm. in diameter, and the conducting tissue 0-2 mm.; the caps of fibres extend
to the epidermis on each side, but separated from it by a single row of thick-walled
cells; the gap between the upper and lower bundles is filled by thick-walled cells
containing starch. There are two smaller veins similar in arrangement to the
midrib; the marginal vein is capped with fibres on the outside only. The peculiar
feature of this species is that there are no columnar stereides, their place being
taken by massive sclereides of extraordinary form in both palisade and medullary
region (Text-figs. 9-13 and 16).

H. undulata R.Br. Leaf 0°34 mm. thick, epidermis 0:03 mm., cuticle 0:01 mm.
Palisade in one row 0:08 mm. deep. Central region a rather loose network 0-16
mm. across. Middle vein 0:44 mm. and conducting tissue 0-16 mm.; there are
besides a number of sub-equal veins all lightly capped with fibres; there are no
columns but many irregular stereides in the medullary region some of which
penetrate the palisade.

H. petiolaris Meiss. Leaf 0:36 mm. thick, epidermis strongly ridged, 0:06 mm.,
cuticle very thick, 0:04 mm., vestibule bell-shaped, the surrounding cells rising
above the leaf surface, 0:06 mm. wide. Palisade in one row, 0:08 mm. on the
ventral and 0:06 mm. on the dorsal side. Central region 0:14 mm. across. The

272 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, i,

main vein is 0-76 mm. in diameter, and the conducting bundle 0-36 mm., the fibres
completely surround the vessels. There are no columns or detached sclereides.

H. neurophylla Meiss. Leaf 0:32 mm. thick; epidermis ridged, 0:04 mm.,
cuticle 0:015 mm., the surrounding cells projecting above the leaf surface. Palisade
one row 0:1 mm. deep. Medullary region 0-12 mm. wide, the cells closely packed.
Principal vein 0:46 mm. the conducting vessels being 0:18 mm. ‘There are no
columnar or detached sclereides.

H. cucullata R.Br. In this species the leaves are wide, rounded, concave ana
arranged horizontally on the stem, but the structure is centric. Leaf 0°56 mm.
thick, epidermis 0:08 mm. and cuticle 0°04 mm. In specimens from the Melbourne
Botanic Gardens the surface is ridged, but in those from the Sydney Gardens it
is quite smooth. The vestibule is bell-shaped but in a few instances it is constricted
in the middle (Text-figs. 6-7) much as in Dasylirion /filifolius, figured by
Haberlandt.“ The palisade is one-rowed and is 0:08 mm. deep. The central
region is 0:2 mm., the cells very full of starch. Main vein 0:48 mm., and conducting
tissue 0-12 mm. The cap of fibres extends to the epidermis on each side, and is
only separated from it by a layer of thick-walled cells; on each side of the
conducting bundle and between the ends of the bundles of fibres there are
about twenty thick-walled cells full of starch. The specimens from Melbourne
Gardens have many irregular columnar sclereides sometimes penetrating deeply
into the medullary region, especially where the section passed through an injury
to the leaf. The specimens from the Sydney Gardens have no columns, but in
the central region there are occasional detached sclereides.

H. conchifolia and H. Victoriae. These are generally considered to be
varieties of H. cucullata; the structure is exactly similar.

H. ambigua Meiss. (Plate xxiii, fig. 2). The leaf is 0:'56 mm. thick, epidermis
0:06 mm., ridged, cuticle 0:04 mm., vestibule bell-shaped. Palisade in two rows,
*0-14 mm. on the ventral and 0:12 mm. on the dorsal side. Central region 0:12 mm.
across, the network rather close. The principal vein is 0-5 mm. across and the
conducting tissue 0:14 mm.; the whole bundle, as in H. petiolaris, is entirely
enclosed by sclerenchyma. The columns are few, they branch in an extraordinary
manner, sending out horizontal arms between the two rows of palisade and from
these upright processes grow between the cells of the palisade like candelabra
(Text-figs. 26 and 27).

H. plurinervia F.v.M. Leaf 0:32 mm., epidermis 0:02 mm. and cuticle 0-01 mm.
Palisade in one row 0:06 mm. deep. Central region 0:06 mm. Vein 0:38 mm., the
conducting tissue being 0:16 mm. There are no columnar or detached sclereides.

H. dactyloides Cay. I have described this species in These ProckeEDINGS,®?
and there is nothing to add except to correct the mistake I made in calling the
persistent bases of the two-armed hairs capitate hairs. A noteworthy point is the
extraordinary massive sclereides, figured in the paper.

H. oleifolia R.Br. Leaf 0:34 mm. thick, epidermis 0:04 mm. and cuticle
0:01 mm., vestibule wide, bell-shaped, 0:06 mm. wide. Palisade one row, 0:06 mm.
deep. Central region 1 mm. across. Vein 0:36 mm. in diameter, conducting tissue
0:24 mm. This species is chiefly remarkable for the absence of columns and the
very small strands of sclerenchyma. It is, like H. lawrina, very mesophytic in
type.

H. stenocarpa R.Br. Leaf 0-38 mm. thick; epidermis 0:06 mm., cuticle 0:03 mm.
Palisade in one row, 0:01 mm. deep; central region 0:1 mm. across. The chief
feature is the great development of fibrous sclerenchyma round the midrib. It

BY A. G. HAMILTON. ; 273

is 0:84 mm. vertically and extends 1:3 mm. horizontally, the conducting tissue
being 0:28 mm. There are no columnar stereides.

H. varia R.Br. The leaf is 0:54 mm. thick; epidermis 0:06 mm., cuticle
0:02 mm., the vestibule narrow bell-shaped with the cells surrounding the orifice
projecting to a height of 0-06 mm. above the surface of the leaf. Palisade in one
row, 0:12 mm. deep. Medullary region 0-1 mm. across. Centre vein 0:42 mm. in
diameter, the conducting tissue being 0-22 mm. There are many columns which
spread out at the apex and base so as to form sheets under the epidermis and
above the central region.

The variety H. intermedia is identical in structure with H. varia.

H. nitida R.Br. Leaf 0-54 mm., epidermis 0:06 mm., cuticle 0:02 mm., vestibule
cylindrical but slightly constricted in the middle so as to resemble a dice box,
0:03 mm. wide. Medullary region 0-1 mm. across. The veins are 0:36 mm. in
diameter, the conducting tissue being 0:28 mm. The columnar stereides are slender
and irregular, the bases extending into the central region.

I have examined an undescribed species in the Sydney National Herbarium,
collected by the late J. H. Maiden at Esperance, W.A. It has the ordinary type of
structure, the leaf measurements being: Leaf 0-5 mm., epidermis 0:04 mm., cuticle
0:02 mm., palisade one row, 0:18 mm., medullary region 0-12 mm., rather loosely
packed, veins 0:35 mm., and conducting tissue 0:14 mm. The columnar stereides
are slender and not spreading at the base or apex.

I have not been able to procure specimens of the following species:—
H. stenophylla, H. trineura, H. stenocarpoides, H. cinerea, H. loranthifolia, H.
smilacifolia and H. carina. There are also a few more recently described species
which I have not seen.

I am much indebted to the late Professor A. A. Lawson, Professor T. G. B.
Osborn, Professor Adamson (who examined all my sections), and Dr. J. McLuckie
for valuable advice and assistance; to the late Mr. J. H. Maiden for access to the
National Herbarium; to Messrs. L. Rodway, H. Cheel, F. Blakely, C. T. White,
J. Audas, C. A. Gardner, O. H. Sargent, J. Steedman and C. G. Hamilton for
material; and to Mr. F. Booker for photographing illustrative sections, and
assistance in preparing text-figures.

Literature.

Solereder gives a comprehensive bibliography (although he omits the Australian
papers on the subject), but many are theses or dissertations, and no copies could be
traced in the Sydney libraries.

@ BENTHAM, G.—Flora Australiensis.

© CANNON, W. A.—‘“Plant Habits and Habitats in the Arid Portions of South Australia,’
Carnegie Publications, No. 308.

@) CHEEL, E.—Proc. LINN. Soc. N.S.W., 1918, xliii, p. 255.

® HABERLANDT, G.—Physiological Plant Anatomy, English Ed. (1914).

©) HAMILTON, A. G.—Proc. LINN. Soc. N.S.W., 1914, xxxix.

® MouH.u, H.—‘‘Der Spalt6ffnungen auf den Blattern der Proteaceen,’ Acta. Acad. Caes.

Leop. Carol. Nat. Cur., Vol. xvi (1833).
© SoLEREDER, H.—‘‘Systematic Anatomy of the Dicotyledons,”’’ English Ed., 2 Vols. (1908).

EXPLANATION OF PLATES.
Plate xxiii.
Vestibule of stomate showing thickenings in wall of orifice.
Root-like extensions of columns in H. ambigua.
Veins surrounded by cells containing tannin, H. ceratophylla.
Sclerenchyma spreading into epidermis, H. Brownii.

wm CO bo bt

274 XEROPHYTIC STRUCTURE OF LEAF IN AUSTRALIAN PROTEACEAE, i,

Plate xxiv.

Thick-walled cells occupying the mesophyll, H. Browvnii.

2. Section parallel to plane of leaf showing arrangement of palisade and sclereides
round cavity beneath stomate, H. pandanicarpa.

Thin-walled cells of mesophyll, H. clavata.

4. Spongy tissue occupying the mesophyll, H. wndulata.

a

(av)

Plate xxv.

Columnar stereides spreading at apex and base, H. Brownii.

Mesophyll occupied by very thick-walled cells, H. obtusa.

Tannin cells running through centre of mesophyll, H. pandanicarpa.

Mesophyll occupied by elongated cells containing chlorophyll, H. amplexicaulis.

mow be

AUSTRALIAN HESPERIIDAE. Part i.
Nores AND DESCRIPTIONS OF NEW Forms.
By G. A. WaterHousE, D.Sc., B.E., F.E.S.

(Plate xxvi.)
[Read 31st August, 1927.]

Since “The Butterflies of Australia” was published in 1914, further collecting
has resulted in records of many new localities together with several new forms
of skippers; the more important of these are given in the present paper.

Subfamily TRAPEZITINAE.

Watson (P.Z.S., 1893, p. 71) places the species I include in the Trapezitinae in
Section A of his subfamily Pamphilinae, and remarks (p. 69) that they are some-
what aberrant, and in his key (p. 71) puts the Australian genera in a group
distinct from the other genera of his subfamily Pamphilinae. As this group of
Australian genera is confined to Australia, New Guinea and the neighbouring
islands, a new subfamily was proposed for them. In “The Butterflies of Australia”
51 species of this subfamily were placed in ten genera and it was pointed out
(p. 175) that when the life-histories of the species became known, further
distinctions would be found. By careful searching I have found some portion
of the early stages of 20 species and as the result of a study of these, I have
found it necessary to alter the sequence of the genera.

The early stages may be divided into three groups: (@) Larva with head large,
hard, rough, some shade of brown without any conspicuous markings; body some
shade of brown, often with a pinkish tint and somewhat rough; pupa stout, with
the anterior end smooth without any prominent projection. Here are placed the
genera Trapezites, Anisynta, Signeta, Dispar and Toazidia. (bv) Larva with head
large, hard, black, covered with long white hairs; body white, hairy; pupa
without prominent projections at anterior end. Only Mesodina comes here. I
know the life-histories of halyzia and aeluropis, both of which pupate head down-
wards. (c) Larva with head large, hard, pale with a dark brown or black band
almost encircling the head in front; body smooth, pale green or yellowish, outer
skin very transparent; pupa longer than those of (a), cylindrical, tapering
towards posterior end, always with a prominent projection at anterior end, some-
times this projection is long and bifid and varies considerably in the different
species. Hesperilla and Oreisplanus come into this section, which, from their
larval characters, are allied to those Australian species of the genera Taractrocera,
Padraona and Telicota. The only genus of Trapezitinae omitted is Neohesperilla
and of the early stages of its species, I know nothing at present.

From the various characters I would now place the genera in the following
order: Trapezites, Dispar, Signeta, Anisynta, Toxidia, Mesodina, Neohesperilta,

K

276 AUSTRALIAN HESPERIIDAE, i,

Motasingha, Oreisplanus and Hesperilla. The larval and pupal characters confirm
the slight imaginal characters used in separating Toxidia and Hesperilla.

It is very difficult to draw up satisfactory descriptions of the species in the
Trapezitinae, as though they appear very distinct in the cabinet, the markings
above are very similar and similarly placed, so the descriptions of the upper-
side are very much the same. The underside of the hindwing always shows the
chief distinctions and the apex of the forewing beneath is always coloured in the
same manner as the hindwing beneath.

TRAPEZITES ELIENA Hewitson.

Hesperia eliena Hew., Desc. 100 new Hesp., p. 32, 1868 (Moreton Bay) ;
Trap. eliena Herr-Schff., Stett. Hnt. Zeit., 1869, p. 80, pl. iii, fig. 13 (Brisbane) ;
Telesto caecilius Plotz, Stett. Ent. Zeit., 1884, p. 380 (India in error); Trap.
iacchus Meyr. and Lower (nec Fabr.) Trans. Roy. Soc. 8S. Aust., 1902, p. 87;
Trap. eliena Waterh. and Lyell, Butt. Aust., p. 176, figs. 604, 605, 1914.

From the large number of specimens available, I can distinguish two good
subspecies.

TRAPEZITES ELIENA ELIENA Hewitson.

I possess many specimens from the type locality, Brisbane, and also from
Kuranda, Westwood and Gayndah in Queensland, Richmond River, Sydney, and
Blue Mountains in New South Wales. In these the general colour of the apex
of the forewing and the whole of the hindwing beneath is orange brown in the
males and very slightly paler in the females. The spots on the hindwing beneath
are sometimes reduced to small blark dots without whitish centres, but the discal
row are never absent; when reduction takes place the central spot is also reduced
in size.

TRAPEZITES ELIENA MONOCYCLA Lower.

Trans. Roy. Soc. S. Aust., 1911, p. 1389; Trap. iacchus And. and Spry (nec
Fabr.) Vict. Butt., p. 115, figured, 1893.

I have examined the type from Mt. Gambier, S. Aust., and also the specimens
from Victoria in the National Museum, Melbourne and in Mr. Lyell’s collection
and consider this a southern subspecies, characterized by the apex of the fore-
wing and the hindwing beneath being almost yellow. The central white spot of
the hindwing beneath is always large and the discal row of four spots reduced
in most cases to mere dots, sometimes absent as in Lower’s type. I have 6 ¢ and
4 9° from Victoria (Gisborne, Fernshaw and Mordialloc) all of which can be
recognized from northern specimens by the underside, though in no case have the
discal spots entirely disappeared. I would restrict the name monocycla to
S. Australia (type) and Victoria. I have not seen a specimen of eliena from
Tasmania.

TRAPEZITES IACCHOIDES Waterhouse.
This rare species has been taken at Barrington Tops in December by Messrs.
Goldfinch and Burns. The specimens differ slightly from Sydney specimens, being
much brighter in colour.

TRAPEZITES PHIGALIOIDES Waterhouse.
I have taken this species at French’s Forest, near Sydney, in October.

BY G. A. WATERHOUSE. 277

SIGNETA TYMBOPHORA Meyr. and Lower.
This very local species was sent to me by Mr. EH. J. Dumigan from Bunya
Mts., Qld. (Jan., 1926). This is a new record for Queensland, the species hitherto
having been found only in the Illawarra District, New South Wales.

ANISYNTA SPHENOSEMA Meyr. and Lower.

I found several larvae in King’s Park, Perth, W. Aust., in September and
succeeded in rearing a specimen which emerged in Sydney in March. The larvae
and pupae are very similar to those of Trapezites.

ANISYNTA TILLYARDI Waterhouse and Lyell.

The range of this species has been extended from the type locality (Ebor,
N.S.W.) to Bunya Mts., Qld. (E. J. Dumigan, Jan.), Murrurundi (Febr.), and
Barrington Tops, N.S.W. (Jan., G.A.W.).

ANISYNTA TASMANICA Miskin.
Additional localities in New South Wales at which I have caught this species
are Stanwell Park (near sea level, Mar.), Blackheath (Nov.) and Barrington Tops
(Jan. and Febr.).

TOXIDIA MALINDEVA Lower.
Taken at Westwood, Qld. by Mr. A. N. Burns in September and October.

TOXIDIA CRYPSIGRAMMA Meyr. and Lower. (Plate xxvi, fig. 20, di, 19, 9.)

Mr. A. N. Burns has taken this species at Westwood, Qld., in February,
September and October, including a single specimen of the undescribed female.

Q. ABOVE. Forewing dull brown; two small subapical elongate spots, whitish;
discal spots in areas 2 and 3 and a small discal spot above vein 1, whitish; cilia
grey-brown. Hindwing grey-brown; cilia greyish.

BENEATH. Forewing pale grey-brown at apex, brown in cell and on disc,
greyish near tornus, spots as above, but fainter; cilia greyish, slightly darker at
terminations of veins. Hindwing pale grey-brown; cilia as in forewing.

This specimen was taken at Westwood in August, and is in Coll. Burns.
This species is easily distinguished from the larger T. peroni and T. malindeva,
both of which have a spot in cell of forewing and have the hindwing marked
beneath. It is nearest 7. sexguttata, which is a narrower winged species and
has the hindwing beneath a different shade of brown without markings. Males
from Westwood as well as the two from the type locality (Herberton) are larger
than those Mr. R. Illidge has sent me from Brisbane.

TOXIDIA SEXGUTTATA Herrich-Schaeffer.

Mr. L. Franzen has given me a female from Palmwoods; S. Queensland (Febr.),
which is much darker than the five I have from N. Queensland.

MorasINGcHA MONTICOLA OlIJiff.
New localities are near Gisborne, Vict. (Lyell, Mar.), and Jenolan Caves, N.S.W.
(L. H. Moss-Robinson, Febr.). Though I have visited Mt. Kosciusko each month
from December to March, I have never seen this species there, probably as I
collected only above 5,000 ft. Prof. L. Harrison has given me two specimens
caught at 4,000 ft. in February.

278 AUSTRALIAN HESPERIIDAE, 1,

MoTaASINGHA DOMINULA Pl6tz.

The locality given by Pl6tz is Tasmania, but his measurement is larger
than any Tasmanian specimen I have and the unpublished coloured figure by
Ploétz agrees better with New South Wales specimens than Tasmanian. Should
Tasmania be correct for dominula, Meyrick’s name drachmophora must be used
for the mainland specimens, which I have taken at Mt. Kosciusko in February
and March and Barrington Tops in January and February.

HESPERILLA ANDERSONI Kirby.
A single male was taken by me at Murrurundi in February.

HESPERILLA IDOTHEA Miskin.
I have caught this handsome species at Barrington Tops in January and
February.

HESPERILLA DONNYSA Hewitson.

Desc. 100 new Hesp., p. 39, 1868; Hewitson, Hxotic Butterflies, v, Cyclopides
and Hesperilla, fig. 7, 1874. 2

“Upperside. Male—Anterior wing rufous-brown, with six transparent yellow
spots; one near the middle of the costal margin, two central beyond the middle,
and three before the apex; a small yellow spot near the inner margin, and the
black line indicative of the male. Posterior wing dark brown, with the centre
orange. |

“Underside grey. Anterior wing with the basal half dark brown; a ray of
yellow from the base to the first transparent spot; the inner margin yellow
in the place of the small yellow spot of the upperside. Posterior wing with a spot
before the middle, followed by a straight band of small brown spots.

“Hemale without the small yellow spot of the anterior wing and the central
orange of the posterior wing. Exp. 1°/, in. Hab. Australia (Moreton Bay). In
the collection of W. C. Hewitson’’.—Hewitson l.c.

In Exotic Butterflies, Hewitson modifies this description slightly, giving the
orange spot of the hindwing in the male as trifid, and the hindwing beneath “with a
minute black spot before the middle, followed by a straight band of several similar
spots’. The upperside of the male only is figured and a slightly increased size is
given.

I have 127 specimens of H. donnyse in my collection, but unfortunately do not
possess a specimen from the type locality, Brisbane. When this series is examined
in the cabinet, certain well marked differences are seen, the orange spot of the
hindwing in Tasmanian specimens is always larger and brighter than in specimens
from Australia. A small series from Altona Bay, Vict., given me by the late
F. P. Spry many years ago, are all very pale and approach another series of
specimens bred by me last year from Geraldton, W.A. These three series constitute
the three new races I describe. Regarding South Australian specimens, I was
inclined to consider them as H. cyclospila, Meyr. and Lower, 1902, when I
examined the type in the late O. B. Lower’s collection in Adelaide last year. Mr.
N. B. Tindale has at my request made a further examination of the specimens in
the Lower collection under the name cyclospila and considers they constitute a
species distinct from donnysa.

Some confusion has arisen with regard to the name cyclospila, as fig. 632 in
“The Butterflies of Australia” is certainly not cyclospila, though Lower identified

BY G. A. WATERHOUSE. 279

it as such. Meyrick and Lower were quite right in placing cyclospila near donnysa,
and it may possibly be a subspecies of it, whilst the specimen given as fig. 632 is
certainly closer to chrysotricha M. and L. H. donnysa has a more pointed fore-
wing (especially in the male) than chrysotricha and its allies.

The four races of H. donnysa may be distinguished as follows:—

HESPERILLA DONNYSA DONNYSA Hewitson. (Plate xxvi, figs. 1, 2, 6.)

dg. ABOVE. Forewing dark brown, with six hyaline yellowish spots, one large
in cell, three small subapical and two beyond the sexmark, an opaque yellow spot
above middle of vein 1; base faintly dusted yellowish. Hindwing dark brown, with
a large dull orange spot divided by the veins.

BENEATH. Forewing with apex grey suffused lilacine, cell broadly pale yellow,
transparent spots as above, dorsum whitish. Hindwing grey suffused lilacine, a
small spot in cell and a discal series of six spots (five almost in a straight line),
brown.

9. ABOVE and BENEATH as in male, but all spots larger.

I would restrict the typical race to Brisbane, New South Wales and Eastern
Victoria. In Exotic Butterflies, Hewitson gives a coloured figure of the upperside
of the male. In “Butterflies of Australia’, figs. 633 and 634 show this form.

I have 24 g, 14 9 from N.S.W., and 12 ¢, 13 9 from Victoria. In two males
there are only two subapical spots and in seven males no opaque spot on the fore-
wing above, in three females there is a second smaller opaque spot above the other.
Beneath the general colour varies somewhat. On the hindwing beneath, the six
discal spots are not always present, in 4 4 7 @ the cell spot is white centred and
in three other males and two other females some of the discal spots are white
centred, this occurs more frequently in Victorian specimens.

HESPERILLA DONNYSA FLAVESCENS, n. subsp. (Plate xxvi, figs. 17, 18.)

¢. ABOVE. Forewing pale brown, almost wholly covered with yellow scales
so that the six hyaline spots are not so transparent as in the type form; two small
opaque spots above vein 1. Hindwing pale brown, dorsum covered with yellow
scales, central spot yellow orange.

BENEATH. Apex of forewing and hindwing yellowish grey, cell spot and discal
spots slightly white centred.

@. ABOVE aS in male with the three subapical spots much longer and a fourth
below them; spot above vein 1 large and a smaller one above it. Hindwing with
an obscure spot in cell and two beyond the large central yellow spot.

BENEATH as in male, all spots of hindwing white centred.

I have four specimens of each sex and there are others in the National
Museum, Melbourne. One male and two females have a fourth subapical spot.

Locality: Altona Bay, Victoria, Mar., Apr. and Nov. (F. P. Spry).

HESPERILLA DONNYSA AURANTIA, n. subsp. (Plate xxvi, figs. 5, 21, 22.)

This form can be recognized by the large bright orange spot of the hindwing
above, in most cases the cell spot of the forewing is larger and often in the
male and usually in the female there are two opaque spots above vein 1, some-
times as in fig. 21 these two spots coalesce, rarely the subapical spots are connected
by two small spots to the discal spots, thus forming a complete band from vein 1 to
costa, broken only by the dark veins. Sometimes there are two obscure pale spots
beyond the central spot of the hindwing above, in the female.

280 AUSTRALIAN HESPERIIDAE, i,

On the hindwing beneath the spots are larger but not so well defined. In
five males and one female the cell spot is white centred and in a further one male
and five females, one or more of the discal spots are centred white as well.

I possess 22 ¢, 13 2 from Tasmania from the following localities: Haglehawk
Neck, 2 Febr., 1910 (holotype d, fig. 22, allotype 9, fig. 21, and two paratype males) ;
Brunei Is., Dec.; Mt. Wellington, Jan., Febr.; Mt. Magnet, Jan.; Strahan, Febr.;
Queenstown.

HESPERILLA DONNYSA GALENA, n. Subsp. (Plate xxvi, figs. 9, 10, 138, 14.)

¢. ABOVE. Forewing brown, basal half covered with yellowish scales; spot in
cell, large, subapical spots usually four, two discal spots beyond sexmark, all
transparent yellow; distad of cell spot a large black streak; sexmark irregular
grey; cilia yellowish. Hindwing brown; dorsum covered with yellowish scales; a
large central trifid spot, yellow; cilia yellowish.

BENEATH. Forewing with costa and apical third grey; transparent spots as
above; a broad yellowish streak in cell; two yellowish spots above middle of
vein 1; rest of wing black. Hindwing grey; central spot and discal series of six
spots, silvery white, ringed with black.

9. ABOVE. Forewing as in male, but spots larger; a large opaque spot above
middle of vein 1 and a smaller one above it, yellow. Hindwing as in male, central
patch, larger and brighter.

BENEATH as in male; spots of hindwing beneath usually larger.

This very distinct subspecies is described from holotype 4, allotype 9, and
9 g, 7 2 paratypes, which I bred from larvae and pupae found at Geraldton, W.A.,
in September, 1926. All but two males and one female (with three) have four
subapical spots on the forewing and two males and one female have still another
spot making almost a complete band from vein 1 to costa. Some males have
opaque spot or spots above the middle of vein 1 of the forewing and six males and
six females have the spots of hindwing beneath, silvery white. The sexmark
differs in colour from the other races. i

HESPERILLA CHRYSOTRICHA Meyrick and Lower.

This species is somewhat allied to H. donnysa, but the shape of the forewing
is different. It is a much stouter-built insect. In all its subspecies the transparent
spots of the forewing are paler than in H. donnysa. At one time I thought that
-it represented H. donnysa in W. Australia, but the male genitalia are different.
I now recognize three subspecies, to one of which I had applied the name
H. cyclospila in error.

HESPERILLA CHRYSOTRICHA CHRYSOTRICHA Meyrick and Lower.

Tozidia chrysotricha Meyr. and Lower, Trans. Roy. Soc. S. Aust., 1902, xxvi,
p59:
The typical form from W. Australia has the apex of the forewing and the hind-
wing beneath dull red brown and the spots of hindwing beneath nearly circular
and centred silvery white.

HESPERILLA CHRYSOTRICHA LEUCOSPILA, n. subsp. (Plate xxvi, figs. 25-28.)

dg. ABOVE. Forewing brown, a small spot at end of cell, yellow hyaline; three
small transverse subapical dots, pale yellow hyaline; a small discal spot in area 3,
pale yellow hyaline; a narrow irregular discal sexmark from before vein 1 to

BY G. A. WATERHOUSE. 281

beyond vein 8; cilia grey brown. Hindwing brown; central patch, deep yellow;
cilia grey.

BENEATH. Forewing brown; apex broadly brown; base of cell pale yellow;
hyaline spots and dots as above; a large discal spot in area 2 and a divided spot
in area 1 whitish; dorsum towards tornus yellowish. Hindwing brown; a spot in
cell, dark brown centred silvery white; a series of discal spots, that of area 1,
small, brown centred white, those in 2, 3 and 5 elongate, white bordered brown,
two dark brown dots in area 4.

%. ABOVE. Forewing brown; cell spot and subapical dots as in male; a large
discal spot in area 3, a minute one in area 2, pale yellow hyaline; a discal spot
above middle of vein 1 yellow; cilia grey brown. Hindwing as in male.

BENEATH. Forewing as in male; subapical spots larger. Hindwing as in male,
white spots much larger; two dots in area 4 better defined.

Locality: Inverloch, Victoria. Holotype dg, allotype 9, paratype ¢ in National
Museum, Melbourne, paratype ¢ in collection Waterhouse. Besides these four
specimens, I have a female from Fernshaw, Victoria, and the South Australian
Museum has a defective female labelled Sheringa, Port Lincoln district, 12:10:89.

This subspecies differs from the Western Australian subspecies, in being
smaller. The markings above, particularly on the hindwing, are not so bright
and the general colour is brown and not red brown. The spots of the hindwing
beneath are twice the length of those of the type race. This subspecies must be

very rare or very local. The figures are of the upper and undersides of the
holotype and allotype.

HESPERILLA CHRYSOTRICHA PLEBEIA, n. Subsp. (Plate xxvi, figs. 23, 24.)

H. chrysotricha cyclospila, Waterh. and Lyell (nec. Meyr. and Lower), Butt.
Aust., p. 188, fig. 632, 1914.

6. ABOVE. Forewing brown; a small spot in end of cell, yellow, hyaline;
three transverse subapical dots, pale yellow, hyaline; a discal spot in area 3 and a
much smaller discal spot in upper edge of area 2, pale yellow, hyaline; cilia grey
brown; a narrow discal sexmark. Hindwing brown; a small central patch, yellow;
cilia grey brown.

BENEATH. Forewing brown; apex broadly dull brown; base of cell yellow;
hyaline spots and dots as above, with that in area 2 much larger; dorsum towards
tornus, whitish. Hindwing brown; a small spot in cell, dark brown centred silvery
white; a series of small discal spots, dark brown, those in areas 1, 2, 3, and 5,
centred silvery white.

Locality: Bridport, Tasmania, two males in December. Holotype in Coll. Lyell
and paratype in Coll. Waterhouse. These two males were identified by Lower as
H. cyclospila, but I have recently examined his type in Adelaide and find that they
do not agree with it. They are smaller than the Victorian race and also paler.
The female is unknown.

HESPERILLA CRYPSARGYRA Meyrick.
This species is nearest to Hesp. picta, but the apex of the forewing is not
quite so pointed. The discovery by myself of a fine large and beautifully marked
race at Barrington Tops extends its range considerably.

HESPERILLA CRYPSARGYRA CRYPSARGYRA Meyr. (Plate xxvi, figs. 3, 4, 7, 8.)

Telesto crypsargyra Meyr., Proc. Linn. Soc. N.S.W., 1887, p. 829; Waterh. and
Lyell, Butt. Aust., p. 186, figs. 600, 601.

282 AUSTRALIAN HESPERIIDAE, i,

On the forewing of the male of this subspecies there is rarely more than
one discal hyaline spot and rarely traces of subterminal streaks above. On the
female these spots are always much larger. The type locality is Blackheath,
N.S.W. (3,500 ft)., and I have caught it as well at Katoomba, Wentworth Falls,
and Woodford (2,000 ft.), all in the Blue Mts.

HESPERILLA CRYPSARGYRA HOPSONI, n. subsp. (Plate xxvi, figs. 11, 12, 15, 16.)

This is a much finer and larger race, almost equalling in size Hesp. picta.
The spots of the upperside in both sexes are always larger and deeper in colour.
On the underside of the forewing in both sexes the general colour is brown
rather than red brown, the spots are orange rather than yellow and there is a
broad orange streak along the upper edge of cell; on the hindwing the silvery
markings are much larger and the veins are orange; the costa towards base is
broadly orange.

I have named this fine form after my friend, Mr. J. Hopson, who has done so
much to help entomologists to collect the treasures of Barrington Tops. Figure 11
is of a male bred in Sydney in October from an egg laid at Barrington Tops in
February and placed on my plants of Gahnia growing in Sydney. Besides the
holotype ¢ and allotype 2 in my collection, I have a number of paratypes of both

25

sexes caught and bred from the same locality in January and February. The
foodplant is a species of Gahnia much coarser and paler in colour than the
narrow-leaved species on which the typical race feeds in the Blue Mts.

Chief References to Australian Hesperiidae.

Lower, O. B., 1911.—Revision of the Australian Hesperiadae, Trans. Roy. Soc. S. Aust.,
XxxV, DD. 112-172.

Meyrick and Lower, 1902.—Revision of the Australian Hesperiadae, Trans. Roy. Soe.
S. Aust., XXvi, pp. 38-129.
MIsKIN, W. H., 1891.—Synonymical Catalogue of the Butterflies of Australia. Annals

of the Queensland Museum. No. 1.
WATERHOUSE and Lyreuu.—The Butterflies of Australia, pp. 172-224. All Australian
species figured.

EXPLANATION OF PLATE XXvVI.

Hesp. donnysa donnysa, 2, Blue Mts., N.S.W., November.

Hesp. donnysa donnysa, 3, Blue Mts., N.S.W., November.

Hesp. crypsargyra crypsargyra, 3, Blue Mts., November.

Hesp. crypsargyra crypsargyra, 2, Blue Mts., November.

Hesp. donnysa aurantia, 3, Mt. Wellington, Tas.

Hesp. donnysa donnysa, 3, Sydney, N.S.W., October.

Hesp. crypsargyra crypsargyra, 3, Blue Mts., November.

Hesp. crypsargyra crypsargyra, 2°, Blue Mts., November.

9. Hesp. donnysa galena, 9, Geraldton, W.A., September. Allotype 9.

10. Hesp. donnysa galena, 3, Geraldton, W.A., September. Paratype ¢.

11. Hesp. crypsargyra hopsoni, 3, Barrington Tops, N.S.W., October. Holotype ¢.
12. Hesp. crypsargyra hopsoni, 9, Barrington Tops, N.S.W., January. Paratype 9°.
13. Hesp. donnysa galena, 9, Geraldton, W.A., September. Paratype, 9.

14. Hesp. donnysa galena, 3, Geraldton, W.A., September. Holotype ¢.

15. Hesp. crypsargyra hopsoni, 3, Barrington Tops, N.S.W., February. Paratype @.
16. Hesp. crypsargyra hopsoni, 2, Barrington Tops, N.S.W., February. Allotype 9.
17. Hesp. donnysa flavescens, 2, Altona Bay, Vict., April. Allotype 9.

18. Hesp. donnysa flavescens, ¢, Altona Bay, Vict., November. Holotype ¢.

13. Tozidia crypsigramma, 2, Westwood, Qld., August. Holotype 9.

20. Toxidia crypsigramma, ¢, Westwood, Qld., October.

21. Hesp. donnysa aurantia, 9, Eaglehawk Neck, Tas., February. Allotype 9.

22. Hesp. donnysa aurantia, 3, Eaglehawk Neck, Tas., February. Holotype ¢.

23. Hesp. chrysotricha plebeia, ¢, Bridport, Tas., December. Holotype ¢.

24. Hesp. chrysotricha plebeia, 3, Bridport, Tas., December. Paratype ¢.

“1m O' & co bo

oo

BY G. A. WATERHOUSE. 283

25. Hesp. chrysotricha leucospila, 9, Inverloch, Vict. Allotype @.
26. Hesp. chrysotricha leucospila, 2, Inverloch, Vict. Allotype 9°.
27. Hesp. chrysotricha leucospila, 3, Inverloch, Vict. Holotype ¢.
28. Hesp. chrysotricha leucospila, ¢, Inverloch, Vict. Holotype ¢.

All figures from retouched photographs. Fig. 25 is the underside of fig. 26 and
fig. 28 the underside of fig. 27. Photographs of figs. 1-24 by Miss A. G. Burns, and
figs. 25-28 by Mr. J. Clark.

NOTES ON AUSTRALIAN MOSQUITOES (DIPTERA, CULICIDAE).

Part ii. THE ZOOGEOGRAPHY OF THE SUBGENUS OCHLEROTATUS,
witH NOTES ON THE SPECIES..

By I. M. Macxerrras, M.B., Ch.M., B.Sc.,
Linnean Macleay Fellow of the Society in Zoology.

(Hight Text-figures. )
[Read 27th July, 1927.]

Edwards in 1924 recognized seventeen valid species of this subgenus of
Aedes from the mainland and Tasmania, later (1926) raising the number to
twenty. In the present paper a further new species is added while two names are
reduced to synonymy, bringing the total down to nineteen. The total represen-
tation of the subgenus in the Australasian region is twenty-two species.

Two distinct faunal elements are to be recognized (Edwards, 1924):

A. Females with well developed lower mesepimeral bristles; male hypopygium with
well developed apical and reduced basal lobes to the side-piece, appendage of
the claspette well developed. Affinities Holarctic. 14 species.

B. Females without lower mesepimeral bristles; male hypopygium with well
developed basal and reduced apical lobes to the side-piece, appendage of the
claspette reduced, usually bristle like. Affinities Neotropical. 8 species.

It is with the origins of these two groups and the explanation of the apparent
anomalies of their distribution that we are here primarily concerned.

The same author (19226) epitomizes the distribution of the subgenus in the
following terms: “This subgenus includes the great majority of the Palaearctic
and Nearctic species of Aedes, and attains its maximum development in the north
temperate parts of these regions; it is almost or quite unrepresented in the
Ethiopian and Oriental regions, but appears again with numerous representatives
in southern Australia, and one or two in New Zealand. Many species also occur ~
in the Neotropical region, and it is possible, therefore, that Australia was colonized
by way of South America, especially as some of the South American, New
Zealand and Australian species show rather marked affinities. We may perhaps
assume from this that the subgenus is an old-established one’.

The Oriental fauna as known at present, and it must be remembered that
this region has been well collected, is entirely deficient in Ochlerotatus, the only
species recorded (Edwards, 1922a) being a Palaearctic invader in the Punjab
and an Australian invader (A. vigilar Sk.) in the south-east. There is no reason
whatever to suppose that the subgenus once flourished but has now become
extinct. The barrier was probably largely, if not entirely, a climatic one.

There is, then, definite evidence for the belief that Australia was colonized
from South America. This evidence is greatly strengthened in the case of
Group A by the local distribution in eastern Australia, a line of investigation

BY I. M. MACKERRAS. 285

which also clearly indicates that the two groups followed different routes and
entered this country from different directions.

In Group A, nine species are strictly “Antarctic” in distribution, i.e.,
dominant in Tasmania, while those occurring also in New South Wales are
restricted to the highlands of the Divide with extensions to the coast in spring.
This distribution is particularly well exemplified by <A. flavifrons Sk. and
A. camptorhynchus Thoms. A. nivalis Edw., on the other hand, is a Kosciusko-
Barrington Tops species which apparently misses the Blue Mountains and is
rare in Tasmania, a somewhat different, but equally typical ‘‘Antarctic”
distribution. The other five species require brief individual mention.
A. stricklandi Edw. is southern and western, while A. burpengaryensis Theo. is
known from South Queensland and is represented in Bass Strait by A. clelandi
Tayl. and in New South Wales by a form intermediate between these two, which
was found at Myall Lakes in spring. These species conform to the same genera]
faunistic localization. A. sagax Sk. (including A. wilsoni Tayl.) is distributed
west of the Divide and is of little value in the present discussion, although its
greater prevalence in the more southerly part of its range is what one would
expect in a member of the ‘‘Antarctic”’ group. Finally, A. vittiger Sk. is a summer
species ranging from Townsville, Qld., to Leeton, in southern New South Wales,
a distribution which would conform quite well with a northern origin.

It must be remembered, however, that the limitation of different faunal
elements to particular environments, though very striking, is not absolute. There
is a definite amount of overlapping and similar extensions are met with in
individual species of other groups. The important fact is that the local
distribution of Group A is precisely that of other Diptera which are believed to
have entered Australia from Antarctica.

Group B presents a different problem. Its Neotropical affinities would
suggest an antipodal origin, but the local distribution is unequivocally against
Antarctic radiation. Of the eight species A. imprimens Walk. is found in New
Guinea and the adjacent islands, A. vigilax Sk. is widespread on the east coast
of Australia and extends into the Oriental region as far as Siam, A. normanensis
Tayl. is not known south of Queensland, while A. rubrithorax Macq. and
A. aculeatus Theo. are both coastal species ranging from South Queensland into
New South Wales. All these species point in the clearest manner to an entry
from the north. A. theobaldi Tayl., like A. sagax Sk., is distributed west of the
Divide from Queensland to Victoria and is therefore of little value, though its
greater prevalence in the more northerly part of its range tends to bring it into
line with the other species of its group. Two species, A. antipodeus Edw. and
A. albirostris Macq., are found in New Zealand and are important as indicating
an eastward extension of the line of migration which brought them to Australia.
I would suggest that this group entered Australia from the north by one of the
Gondwanaland arcs of islands which Harrrison (1927) indicated as being very
successful in explaining other similar distributions.

The more general lines of evidence cannot be entered into here, but the
conclusions may be summarized in the following terms. Group A developed in
the temperate parts of the Holarctic region and was unable to extend to the
southward except along the ridge of cooler elevated country on the western side
of the Americas. Keeping to this higher western side, it extended into the
temperate southern part of South America and radiated thence through
Antarctica into southern Australia, where its climatic limitations are very well
shown in the local and seasonal distribution. This migration occurred at a time

286 NOTES ON AUSTRALIAN MOSQUITOES, ii,

when New Zealand was cut off from participation in this element. Group B, on
the other hand, originated in the Neotropical region, and almost certainly in the
warmer, more low-lying eastern portion. Its line of eastward extension was
directly in warm latitudes. Such an extension implies the existence of a
Gondwanaland, or of its equivalent in a chain of islands (Harrison, 1927) linking
South America to northern Australia and extending eastward to New Zealand.
Both these radiations would ante-date the Oriental invasion which is the
dominant element in the Australian mosquito fauna. It will be seen that these
conclusions imply the acceptance of the Wegener or some similar hypothesis.

The early stages of but two species, A. vittiger Sk. and A. vigilar Sk., have
been described, though a number have been bred. The larvae of all the species
are very difficult to find, even in districts where the adults are so numerous as
to be a pest. Even when it is possible to find a few pupae, the larvae frequently
cannot be discovered. In certain instances, however, they appear to be able to lie
hidden in the mud at the bottom of a pool for prolonged periods.

The male hypopygium presents valuable group characters which are, in all
the Australian species, sufficiently well marked to afford easy and rapid deter-
Mination. Such species as A. theobaldi Tayl., however, show the undesirability
of elevating the two groups to separate subgeneric rank. The presence or absence
of lower mesepimeral bristles on the other hand is not quite so constant, since
these structures are absent in the male of A. flavifrons Sk., though well developed
in the female, and they are quite variable in A. rubrithorax Macq., but when
present in this species are always very weak. In A. stricklandi Hdw. they are
weaker than is usual in Group A, but are markedly longer and stouter than in
A. rubrithorar Macq. A more convenient though unnatural grouping may be
made on the presence or absence of white rings on the tarsal segments. This
character also is not completely satisfactory, since two species, A. stricklandi
Edw. and A. luteifemur Edw., which are best included with the unbanded forms,
show traces of white rings on one or two segments; they are, however, never
likely to be confused with those species possessing true banding, since in the
former the rings are only apparent if carefully looked for, while in the latter
they are easily visible to the naked eye. In the following key I have included
for the convenience of local workers all mainland and Tasmanian species which
are inseparable in the female from Ochlerotatus. The most easily accessible and
constant characters have been used and the arrangement of species is consequently
in many instances unnatural.

Key to Australian Species of Ochlerotatus, Banksinella, and Aedimorphus (females).

Species of Aedes with the following female characters: ninth sternite small and
retracted, cerci long; head with numerous narrow curved scales (except A. aculeatus
Theo.) ; claws of at least the fore and mid legs toothed.

ie TarsiloLall less with aistinet white basal) banding wrerieirecistehelens)cie)els)ielieieieleienemetene 2
Tarsi unbanded, or at most with traces of white rings on one or two of the basal
STAaeiil ol HARADA GGodOo onan GOOG D Ono DOMUIn ODO Uo OU COO aU an oUamUmbodo D100 440 10

2) \Seutum (creamy, with striking, black wittaey ec cic lodsiei-uteeeleh teienevela1« «fells (o)eioile vittiger Sk.
fobidbbectnrsVoreurs(oar ie (oyelso Mia agin aiciniocieod Did MOH e500 0100.05.00. 010 COMO CIOMODOODIO.6,0 000 0 3

3. Head and sides of scutum with flat oval yellow scales ............ aculeatus Theo.
Head and isides of scutum? with narrows SCArPeS) Misicrcrenictsteieieiebelis\ oe) es oie) 0) +l oiiel e aitenetotey eis 4

AS Wines entirely, darks SsCalediok beri ee cleleiete toto crelavenciete yen tetetereneielc BAshie lade folel euekevenetee eee 5
Wines withuaLt least some white YSCalesigm.-susl iinet sted oles) -l-letelol-Kel Nekaick-\erensie rs 6

Oo

Larger, darker species; femora mottled with white anteriorly ..................

AA Seine OOOO OOO CE OD DIO CIO OEE OA Oo. B.S Olo.0laL camptorhynchus Thoms.
Smaller, reddish species; femora not mottled with white anteriorly ..........

Panay tae aeks ola Dic eta\e tm tale Mane¥edole te alete (oh oka ehs Ravel s ER SCEMOW eR Metonetaleleh ond to te rubrithorax Macq.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

BY I. M. MACKERRAS. 287

Lower mesepimeral bristles present; pale scales on wings fairly numerous, scutal
integument reddish, wing membrane usually with a dark blotch in middle

DELOWAMCOSUAN merotepsiaicia sh cies: eNovat rea sioner alousisieus She: dsuekevelevevel oy sbeusseh ele one flavifrons Sk.
Lower mesepimeral bristle absent; scutal integument pitchy, wing membrane without
GAT KAM DIOL Wemayee eater ene re trays rereeibed eneremer a iat mac suelo Novetoper es tereticricie, crey a aan ee aterellow chose ah create 7
Wings with numerous large white scales on all veins ..............22c0eeeeeees 8
Wing scales normal in size and shape, white scales relatively few and mainly along
SE Gin IR, coadodoosodgadoonoopEdodeoOD OOD ODO OUGDOO ADO ODO DO OU DONO DOOOSC 9

Pale bands of abdominal tergites nearly straight posteriorly, the dark portions of the
tergites usually conspicuously mottled with pale scales, especially posteriorly
0B COO TOO ROO OLD Cae OEIC HO LRA OIG OI BeOS DiGi CDOT ER RCI ER ACTS ne Pee rena es theobaldi Tayl.

Pale bands of abdominal tergites broad and produced posteriorly into a prominent
triangle which forms a broad vitta on segment 6, only a trace of mottling on
therdark portionsiot ithe) tereitesiia. vaca. fecal cne form eidsvoldensis nov.

Medium sized species; the gold scales of the scutum fine and patchy; abdomen with
broad white basal bands; last hind tarsal segment with a broad white
DELI Se epee creas hore nat ea ote TSR Ae ER Pe rated CaM Pee eealtcner Seelam aNann tetas ea ais vigilax Sk.
Smaller species; the gold scales of the scutum larger, more numerous and tending
to a linear arrangement; abdomen usually with small white basal patches;
last hind tarsal segment without, or with very narrow, basal white ring
RR es omen peera Se as Laie rea eS Sede semaine cane Me LametmraR el she oneneds Co ob etal evene normanensis Tayl.

Wings extensively mottled with light and dark scales ............ stricklandi Edw.
Pale scales, if present on wing, relatively few and only along anterior border .. 11
Scutellum with broad flat white scales ........ Aedimorphus alboscutellatus Theo.
Scucellumenwithwn arr owasCalesmonliysurwa a ciccrlelcrecncielsieiercieicielchelercienchekeneiensienencnenenensh ite 12
Lower mesepimeral bristle absent; scutum with a broad margin of yellow scales

De ee LEENA Uy RCE ene eteee oH aor atcnaL Nt Lae el as ea aR Banksinella lineatopennis Ludl.

At least one strong lower mesepimeral bristle present ..................200000- 13
Scutum with a broad margin of pale scales contrasting strongly with the dark

MENG “seb bos AS aU rep east Manan cite tahrcs ck eit ayle tay Stasi's valdeiret Sees autal cieeartAve tanick Suaeataial caw! arated cetunat rleusronevee 14
SCutumBenole SoMa GORMe yew eres ere a ehel ten euo late aisicsnaskelepeaaia suaususties crs Ryetisheveletaiensnene 15

Numerous large flat scales above and in front of the wing root; abdomen unbanded
Pee R ee alr atietis. Mont sh chee ROMCHAC eenente wate cites Sesto en wile eRe CPE cotatia cay etre semana sttees asters: eile andersoni Edw.
Large scales above the wing root lanceolate and twisted; abdomen conspicuously

asaillives DamGed. sherri ces cireheme each sees on oaltatanat oes scievewete sone sttouauets macleayanus, Nn. Sp.
Hind femora mottled anteriorly with pale scales ................cecceccececes 16
Hind femora uniformly pale anteriorly at least on the basal two-thirds .......... ily’
Mesonotum dark brown to pitchy; mesonotal scales bronzy-brown to creamy white

SRUZE CESS OO ERO CRO NCO OREN cutheo cet Io ero) cli b OTGNGIG/O Gr OICOra CrO/o Oe onc Or Cho TERE eIE oR nG sagax Sk.
Mesonotum reddish; mesonotal scales bright pale golden ........ cunabulanus Edw.
Fore and mid femora with at least some white scales anteriorly ................ 18
Morewandsmidy femora yentirelydarkwanterionlyay ee eee cic ono 19

Abdomen with white scales at bases of tergites only; wings entirely dark scaled
Seo eh ree rate Hac st Misyict Voile corsa ta vanyists) epremere ovenSte eWave tral cee eten a call clianer eine) of ereSeial ees et nivalis Edw.
Abdomen unbanded, fourth and subsequent tergites extensively mottled with dull
creamy-yellow scales; wings pale scaled on distal part of C and IM, ooovdDe?

kak eM one icrereherlerel chops eon alteNela oNciionsisweretcu cuca Peteteaoieteterecheialionenon a etstnis corer sche: luteifemur Hdw.
Venter entirely dark purple scaled .......................- purpureiventris Edw.
Venter) swithe numerous: pale) as) swell ais) darks sscales) ieiccieciete) =e) lel eto eieleleleiiclelele 20

Abdominal tergites unbanded; scutal integument brownish-black ................
5b SOLS bb coro OOF ORG ORCI CLOUE ORCRCROLCH GtiCrn id DCR AIG EEO OG oS OIaIO GaaiciOoloD burpengaryensis Theo.
Abdominal tergites with basal white bands; scutal integument reddish ..........
PNR INN aioe sae ot cet etn cen Lit an Site Gi es) ola de OORT c amtaal Sh a Maureiavimteginnieutsi eile atemepan sriexol tence ke clelandi Tayl.

Notes.—This key applies only to the females. In the male many characters, which

are conspicuous and useful in the female, are ill defined, as is the case with the white
scales of the wings, or absent, as may occur with the lower mesepimeral bristles. Except
for very distinctive species, a careful comparison with authentic females and an
examination of the hypopygium afford the only safe means of identification.

A. nigrithorax Macq. is only to be recognized by hypopygial characters. The

female when discovered will probably fall under caption 15.

288 NOTES ON AUSTRALIAN MOSQUITOES, ii,

The following species, not yet Known from the mainland or Tasmania, are
not dealt with in this paper:—Aedes (Banksinella) brugi Edw., New Guinea;
Aedes (Ochlerotatus) albirostris Macq., New Zealand; Aedes (Ochlerotatus)
antipodeus Edw., New Zealand; Aedes (Ochlerotatus) imprimens Walk., New
Guinea and adjacent islands; Aedes (Aedimorphus) vexans Meig., widely dis-
tributed. The last would run down either to rubrithorax Macq. or to vigilax Sk.
in the key; the specimens before me are too abraded to be sure whether there
are any white scales on the wings or not. From the former it is readily
recognized by its much darker thoracic integument and scales and by its mottled
femora, while from the latter it is separable by its shorter proboscis and by the
indentation of the abdominal bands, which are straight in A. vigilax Sk.

A very distinctive feature of the nivalis series is the violet gloss on the
abdomen and legs. It is, however, not always present, and when present may
occasionally be difficult to detect. It is therefore useless as a key character.

Culex australis Erich. is unrecognizable and must be deleted from the list of
Australian Ochlerotatus. The name is replaced by nivalis Edw.

5 6

7 8

Text-figures 1-8. Side-piece and appendagegy of: 1, A. vittiger Sk.;
2, A. flavifrons Sk.; 3, A. camptorhynchus Thoms.; 4, A. sagax Sk.; 5,
A. theobaldi Tayl.; 6, A. normanensis Tayl.; 7, A. vigilax Sk.; 8, A. rubrithorax
Macq. All drawn to the same scale.

BY I. M. MACKERRAS. 289

For the full synonymy and references up to 1924 see Hdwards (1924).
Subsequent publications on the subgenus and those earlier ones which are
directly mentioned are listed at the end of this paper.

Group A.—AFFINITIES HOLARCTIC.

The species included here fall into two natural sections on hypopygial
characters, those with a sub-basal thumb on the medial aspect of the claspette
(A. vittiger Sk., A. sagax Sk., A. nigrithorax Macq., A. burpengaryensis Theo.;
Text-figs. 1 and 4) and those without such thumb (A. flavifrons Sk., A.
camptorhynchus Thoms., A. andersoni Edw., A. cunabulanus Edw.; Text-figs. 2
and 3). It will be seen that forms with banded and with entirely dark tarsi
occur in both sections. I have, however, used this unnatural but more convenient
arrangement, largely because of the difficulty of placing correctly some of the
species which are only known from the female.

a. Tarsi with conspicuous white rings.

AEDES (OCHLEROTATUS) VITTIGER (Skuse).

One of the most distinct Australian mosquitoes on account of the scutal
adornment. Its hypopygial characters (Text-fig. 1) show its affinities to be with
A. sagax Sk. and its allies rather than with A. flavifrons Sk. and A. camptorhynchus
Thoms. The claspette is forked and the apical lobe of the side-piece forms a
pointed medial projection bearing a strong bristle.

The larva has been described by Cooling (1913) and the larva and pupa
figured by Hill (1925a). The combination of short antennae, long frontal hairs,
3-4 branched subantennal hair, relatively long siphon, the presence of some of the
pecten spines beyond the siphonal tuft, and the single row of comb scales will
separate this from any other Australian larva.

At Hidsvold A. vittiger Sk. breeds in clear or muddy waterholes and is Very
abundant in midsummer in forest country and in the vicinity of the river. It
bites by day and is a source of considerable annoyance owing to the persistence
of its attacks. In the Sydney district it is rare and frequents the sheltered
sandstone gullies.

Distribution.—Coastal Queensland and New South Wales from Townsville to
Sydney; also extends west of the Divide. A summer species, most prevalent in
south Queensland. Now recorded for the first time from the mountains and
from south of Sydney: Barrington Tops, 5,000 feet, January, 1926; Leeton,
December, 1926 (K. McKeown).

AEDES (OCHLEROTATUS) FLAVIFRONS (Skuse).

Culicada vandema Strickland.

The types series of Culex flavifrons Skuse in the Macleay Museum comprises
four specimens, the type male and female which undoubtedly represent one
species, another female which is identical with typical A. vandema Str., and a
third female identical with A. camptorhynchus Thoms. With regard to the types,
the female differs from typical A. vandema Str. in its duller colouration, the
rather fewer pale scales on the wing and the markedly fainter blotch on the
wing membrane. The two latter characters are somewhat variable in the
extensive series of A. vandema Str. before me and the colour difference may
simply be due to age. A careful comparison revealed no character on which the
two could be separated. 'In the male type the pale scales on the wing are very

290 NOTES ON AUSTRALIAN MOSQUITOES, li,

few, while the dark blotch has entirely disappeared; lower mesepimeral bristles
are absent. A male from Sydney in my collection, which can be allotted without
doubt to A. vandema Str. also lacks the lower mesepimeral bristle and differs
from Skuse’s type in its slightly brighter colouration and in the fact that the
dark patch on the wing is present though very faint. The hypopygia of the two
are identical. I am indebted to Dr. EH. W. Ferguson for drawing my attention to
the possibility of this synonymy.

A. flavifrons Sk. is easily distinguished in the female by the presence of lower
mesepimeral bristles, the numerous pale scales on the wings, the mottled femora,
and almost always by the dark patch on the wing membrane. Superficially it
bears some resemblance to A. (F.) occidentalis Sk. The male (Text-fig. 2) has the
apical lobe of the side-piece well developed and projecting distally, while the basal
lobe is rounded and bears a row of five fine hairs directed proximo-medially; the
claspette has a well developed appendage and bears a single small bristle a little
beyond the middle; this appears to be the homologue of the bristle on the medial
division in those species in which the claspette is forked.

This species is abundant in Tasmania and appears in considerable numbers
in the coastal districts of New South Wales in early spring, .where it frequents
the sandstone gullies and brush country in company with A. (F.) alboannulatus
Macq. and A. (F.) occidentalis Sk. It is a day biter.

Distribution.—Victoria, Tasmania and King Island (November and December).
New South Wales: Blue Mountains (type series); Myall Lakes, Aug. 1, 1922; Woy
Woy, Oct. 3, 1925; French’s Forest, Sydney, Aug. 17, 1924; Balmoral (Sydney),
Nov. 26, 1922; Lane Cove (Sydney), May 31, 1927 (B. Bertram); National Park
(Waterfall), Aug. 2, 1925.

AEDES (OCHLEROTATUS) CAMPTORHYNCHUS (Thomson).

To be separated from A. flavifrons Sk. by its darker colour, entirely dark
wing scales, and absence of any blotch on the wing membrane. The male, unlike
that of the previous species, has well developed lower mesepimeral bristles. The
hypopygium (Text-fig. 3) differs in the presence of a stout bristle as well as fine
hairs on the basal lobe of the side-piece and in the absence of any bristle on the
stem of the claspette, the apex of which is differently shaped. The two species
are obviously closely related.

In the Sydney district its habits and season resemble those of A. flavifrons Sk.
It is, however, less common, and its local environment is rather strikingly different
in that it is found in the heath on the more level sandstone ridges rather than in
the gullies (Mackerras, 1926). In Victoria it appears to replace A. vigilax Sk.
as the dominant “bush” mosquito (Hill, 1925b).

Distribution—New South Wales: French’s Forest, Sydney, Aug. 17, 1924;
Greenwich, Sydney, Aug. 15, 1925 (B. Bertram); National Park (Gundamaian),
Aug. 2, 1925; Sutherland, Aug. 7, 1926. Also Western Australia, South Australia,
Victoria (abundant), and Tasmania (abundant from October to March).

b. Tarsi entirely dark, or with only traces of pale rings.

AEDES (OCHLEROTATUS) STRICKLANDI Edwards.

Easily recognized by the numerous broad pale scales scattered over the
wing. In the specimen before me, there are traces of white rings at the bases
of the first and second hind tarsal segments only; these might easily be over-
looked, but are obvious enough under the binocular. There are two fairly strong
lower mesepimeral bristles on one side; the other side is obscured by the legs.

BY I. M. MACKERRAS. 291

Distribution.—Western Australia and Flinders Island, Bass Strait. Additional
locality: Mt. Compass, Oct. 27, 1920 (J. B. Cleland).

AEDES (OCHLEROTATUS) ANDERSONI Edwards.

Edwards, 1926, p. 112, new name for Andersonia tasmaniensis Strickland.

Another striking species, which is easily separated from all except
A. macleayanus n. sp. by the scutal ornamentation. The abdomen is unbanded.
Edwards (1926) describes the male hypopygium as follows: “Lobes of ninth
tergite small, with four to six short bristles. Side-pieces with the basal lobes
large, flattened, prominent; a row of long hairs along the margin, and two
tergally placed bristly spines, one long with slender curved tip, the other short
and straight. Apical lobe well developed but practically bare. Claspers not much
swollen, slightly tapering, apical spine long. Claspettes with long stem without
basal thumb, appendage flattened but not very broad, without angle at base’. This
description agrees with the hypopygium of A. camptorhynchus Thoms. very closely
and shows the uselessness of leg ornamentation in phylogenetic studies.

Distribution.—I have seen a number of females from Victoria (Lower Tarwin,
biting by day, G. F. Hill) and from Tasmania, October to December. Also
recorded by Edwards for January and February in Tasmania.

AEDES (OCHLEROTATUS) MACLEAYANUS, DN. SD.

2. Head with creamy narrow curved scales in the middle, more ochreous ones
laterally, and with ochreous upright forked scales; lateral flat scales mostly
white, a few violet brown. Antennae dark brown, torus brownish ochreous.
Palpi rather long, deep brown. Proboscis entirely deep brown.

Integument of scutum pitchy. There is a broad median area of bronzy-brown
fine scales which widens suddenly behind the middle of the thorax; lateral to this
is a broad creamy zone of larger twisted scales, which are particularly large
above the wing roots; this zone of creamy scales extends medially round the
anterior margin of the scutum; further laterally there is a narrow zone of bronzy-
brown scales at the lateral edge of the scutum. Scutellum and median prescutellar
area with creamy white narrow curved scales. Anterior and posterior pronotal
lobes with fine bronzy scales above and larger creamy more outstanding ones
below. Pleurae brown, with dense flat white scales forming a band across the
upper part; there is also a large white patch on the lower part of the sterno-
pleuron. Three strong lower mesepimeral bristles. Wings entirely dark scaled.

Fore and mid femora mottled anteriorly, the pale scales predominating; hind
femora entirely. white anteriorly except for a narrow apical dorsal zone. Tibiae
and metatarsi mottled anteriorly, largely white behind. Tarsi dark. Ungues of all
legs toothed.

Abdomen covered with deep brown scales with a slight violet gloss, the
tergites basally banded with ochreous scales. The bands are triangular in shape,
with the broad base along the base of the tergite and the apex more or less drawn
out, on the sixth segment extending to its apex. There is a large lateral basal
patch of creamy ochreous scales on the fourth segment and larger white patches
at the sides of the fifth and sixth segments. Seventh with a broad median creamy

stripe and white lateral margins. Venter white scaled, except for small apical___

lateral black patches on a few segments.
Length, 7 mm.

LD fog

292 NOTES ON AUSTRALIAN MOSQUITOES, ii,

Holotype ?: Launceston, Tasmania, Oct. 1, 1916 (F. M. Littler), in the South
Australian Museum.

This is a rather large species which superficially resembles A. andersoni Hdw.,
but differs in the absence of flat scales above the wing roots, the darker thoracic
integument, the unmottled hind femora, and the banded abdomen. The scutal
ornamentation and the leg characters will separate it from A. cunabulanus Edw.

AEDES (OCHLEROTATUS) CUNABULANUS Edwards.

Edwards (1912) first mentions this species as a variation of A. australis Theo.
(i.e. nivalis Edw.), but, when describing it (1924), he stated that it was closer to
A. andersoni Edw. but much smaller in size. The mottling of the anterior surface
of the hind femora will separate it from A. nivalis Edw., while the reddish thoracic
integument and bronzy gold mesonotal scales distinguish it from A. sagaz Sk.
I have seen a single female from Tasmania which I place here with some
hesitation. It is not much smaller than A. andersoni Edw. and resembles
A. nivalis Edw. closely in scutal colouration and scaling, but otherwise fits the
description fairly accurately. The relationships of A. cunabulanus Edw. have been
settled by Edwards’s (1926) description of the hypopygium of the male as
“practically identical in structure with that of A. andersoni, the appendage of the
claspette appearing a little broader”.

Distribution.—Tasmania, November to February. Additional locality: Mount
Arthur, Tasmania, December 28, 1915 (F. M. Littler).

AEDES (OCHLEROTATUS) NIGRITHORAX (Macquart).

Only known from the type male and only identifiable on hypopygial characters.
It is possible that future discoveries may show the name to belong to one of the
species dealt with below. Edwards’s (1924) description,of the hypopygium runs
as follows: “Side-pieces with the basal lobes well developed, hairy; apical lobes
slight, bearing two or three rather stout curved spines. Claspers with the basal
two-thirds somewhat swollen, apical third slender, terminal spine long. Claspettes
with a sub-basal thumb on the inner side bearing a small terminal bristle, some-
what as figured by Cooling for A. vittiger; appendage broad and flat, with a
retrorse angle at base. Lobes of ninth tergite each with about six short bristles”.

Distribution.—Tasmania.

AEDES (OCHLEROTATUS) SAGAX (Skuse).

Culicada wilsoni Taylor.

Edwards (1924) recognizes these as distinct, but Ferguson (1926) gives this
synonymy and I believe it to be correct. I have seen females of both forms and
there is little to separate them. In typical sagax the scutum is covered with
brown scales with small patches of white, and the pleurae are spotted with white,
whereas in wilsoni the mesonotal scales are all or mostly creamy and the pleurae
are more densely scaled. There are, however, intermediate forms. The violet
sheen of the abdomen is of no value and may be present or absent in otherwise
typical sagaz. Taylor (1918) gives a photomicrograph of the male hypopygium of
A. wilsomi Tayl.; it is unfortunately not suitable for detailed study, but does not
appear to show any essential difference from that of A. sagax Sk.

This species is to be recognized by its dark scutal integument and dull scaling,
by the mottled hind femora, and by the banded abdomen.

BY I. M. MACKERRAS. 293

¢. Palpi entirely brown, a little longer than the proboscis, terminal segments
swollen and bearing dense long brown hairs; antennae with long brown plumes;
one lower mesepimeral bristle; otherwise similar to the female. Hypopygium
(Text-fig. 4) with the apical lobe of the side-piece pointed and bearing one small
and two larger bristles, basal lobe very small and covered with fine hairs. Claspette
with a sub-basal thumb bearing a short bristle and with the appendage serrated
along its apico-lateral edge. The resemblance to A. vittiger Sk. and to Edwards’s
descriptions of A. nigrithorar Macq. and A. burpengaryensis Theo. is remarkable.
Allotype ¢: Hidsvold, South Queensland, Oct. 13, 1926 (Mackerras), in the Macleay
Museum, University of Sydney.

Distribution.—Widely distributed in Victoria and New South Wales west of
the Divide. Now recorded for the first time from Queensland: Hidsvold, June 6, 1926,
biting by day (T. L. Bancroft), and October 7-18, 1926, females biting by day and
at dusk, male in tussock near a swamp. Taylor records A. wilsoni as a source of
great annoyance in the Goulburn Valley district, Victoria.

AEDES (OCHLEROTATUS) LUTEIFEMUR Eidwards.

Edwards, 1926, pp. 112-113.

This is rather an inconspicuous species, but is readily recognized by the
entirely pale hind femora, especially by the numerous pale scales towards the apex
of C and R,, and by the unbanded abdomen which is uniformly dark anteriorly,
extensively mottled with ochreous creamy scales in the middle and entirely pale
posteriorly.

Distribution.—Tasmania and Victoria (Edwards).—There are specimens in
the South Australian Museum from Georgetown, Tasmania, November to March,
and from King Island, Bass Strait, November and December, all coll. F. M. Littler.

AEDES (OCHLEROTATUS) NIVALIS Edwards.
Edwards, 1926, p. 112, new name for A. australis Theo. nec. Macq.

. Proboscis and palpi dark scaled, the former with a few white scales. Scutum
and scutellum bright reddish to dark purplish brown; clothed with small bright
pale golden scales, sometimes tending to a linear arrangement, but without any
special ornamentation. A row of three to six lower mesepimeral bristies, usually
the larger number. Legs black, with distinct violet reflections in most specimens;
fore and mid femora with some scattered white scales anteriorly, hind femora
white on basal four-fifths anteriorly, dark apically with some scattered white
scales; no pale scales on tibiae and tarsi. Abdomen deep brown, usually with
violet reflections, unmottled; all segments with basal white bands of variable
extent, sometimes reduced to a single incomplete row of white scales, but never
in my experience entirely absent. Venter white, with large apical lateral and
small basal median dark patches on all segments. Size variable, some specimens
up to § mm., the majority about 5-6 mm. The most characteristic features are the
scutal and abdominal ornamentation, the markings of the venter, the adornment
of the femora, and the violet reflections when present.

This species is exclusively alpine in New South Wales. It was exceedingly
abundant and annoying at Barrington Tops in January and February, biting
viciously at dusk and for an hour or two after; occasional individuals could be
taken biting at almost any time.

Distribution—New South Wales: Kosciusko; Barrington Tops, 5,000 feet,
January and February, 1925. Victoria: In elevated country. Tasmania: Recorded

294 NOTES ON AUSTRALIAN MOSQUITOES, ii,

by Edwards (1926); there is also a single female in the South Australian Museum
labelled ‘‘Tasmania” without further data.

AEDES (OCHLEROTATUS) BURPENGARYENSIS (Theobald).

This and the two following are to be separated from all others with dark
tarsi by the complete absence of pale scales on the anterior surface of the fore and
mid femora. The thoracic integument of A. burpengaryensis Theo. is described
as deep brownish black and the abdomen as unbanded but with white basal lateral
patches to the tergites. The venter has yellowish basal bands to the segments.
A specimen before me from Myall Lakes agrees very well with the description, the
almost metallic violet reflections of the legs being particularly striking. The
tergal markings of the abdomen, however, are different and approach A. clelandi
Tayl.; the second and third tergites are unbanded, the fourth has a very narrow
basal band and there are wider basal bands on the fifth to seventh segments. I
have no doubt that this specimen should be placed with A. burpengaryensis Theo.
rather than A. clelandi Tayl., but it certainly forms a link between the two.

Edwards (1924) gives the following note on the male hypopygium: ‘Chiefly
differs (from A. nigrithorax Macq.) in having the spines on the apical lobe of the
side-piece more slender, and the sub-basal thumb of the claspette rather shorter’.

Distribution.—South Queensland: Burpengary (type locality). New South
Wales: Myall Lakes, Sept. 3, 1922 (A. J. Nicholson).

AEDES (OCHLEROTATUS) CLELANDI (Taylor).

I have examined a paratype and another authentic female in Dr. Ferguson’s
collection and cannot find anything to separate it from A. burpengaryensis Theo.
beyond the bright reddish scutal integument, the conspicuous basal banding of
the abdominal tergites and the more extensively pale scaled venter. The violet
gloss of the legs is slight in these specimens, but both are rather wasted. These
characters are hardly, in my opinion, of specific value. I believe it probable that
A. burpengaryensis Theo. and A. clelandi Tayl. will be found to represent the
extremes of a continuous north-south range of variation, but the evidence is as yet
insufficient and I have decided to let Taylor’s name stand for the present.

Distribution.—Bass Strait: Flinders Island, Nov. 21, 1912 (J. B. Cleland) ;
King Island (A. M. Lea).

AEDES (OCHLEROTATUS) PURPUREIVENTRIS Edwards.
Edwards, 1926, p. 113.
I have not seen this species, which should be easily recognized from
A. burpengaryensis Theo. by the entirely purple scaled venter.
Distribution.—Tasmania.

Group B. AFFINITIES NEOTROPICAL.

Tarsi with white rings.

A. aculeatus Theo. stands out as very distinct, but the other species of this
group are all closely similar. The characters given in the key will usually be
sufficient for their recognition, but the variability of certain species may render
separation difficult in the case of extreme specimens. Indeed, out of the hundred
odd females of A. theobaldi Tayl. and A. normanensis Tayl. before me, there are
one or two which might be placed with almost equal propriety in either species.
Hypopygial characters, however, establish their specific validity.

BY I. M. MACKERRAS. 295

AEDES (OCHLEROTATUS) ACULEATUS (Theobald).

A very well defined but rare species with the head and sides of the scutum
covered with small oval flat yellowish scales. It is the only Australian member
of the subgenus without numerous narrow curved scales on the occiput and its
affinities are somewhat doubtful.

Distribution.—South coastal Queensland and north coastal New South Wales.

AEDES (OCHLEROTATUS) THEOBALDI (Taylor).

The most striking feature of this rather ornate species is the presence of
numerous broad pale scales on all the main veins of the wing. Though somewhat
variable in number and size, these scales are always more numerous and distinctly
broader than in any other species of the group. The scutum is covered with small
densely arranged golden scales mixed with darker ones. The abdominal tergites
are basally banded with creamy scales, usually prominent, but sometimes reduced
to a narrow incomplete fascia, never, however, forming a posterior triangular
prolongation. The dark posterior portions of the tergites show a variable amount
of mottling with pale scales; more marked posteriorly; occasionally the mottling
is practically absent and the abdomen is indistinguishable from that of A.
normanensis Tayl., while in some specimens the whole abdomen is mottled and
the basal bands are ill defined. The femora are extensively mottled anteriorly
with white scales and the last hind tarsal segment is entirely dark or with a
narrow basal white ring.

6. Differs from the female as follows. Palpi brown, slightly longer than the
proboscis, with some creamy mottling beneath which extends round the shaft in
the middle to form an indefinite pale ring; penultimate segment with a narrow
basal white ring; the apical segments slightly swollen and bearing long brown
hairs. Antennae with long silky brown plumes. The abdominal bands are
definite and there is little or no mottling. The white scales on the wing are
scanty and scattered, but are more numerous and broader than in the males of
related species. The side-piece (Text-fig. 5) bears an elongate rather hairy apical
lobe and a very large characteristically shaped basal lobe covered with fine hairs
which form a definite fringe on its medial border. The appendage of the claspette
is better developed than in the other members of the group and forks distally to
form a striking fluke-like structure. A. theobaldi Tayl. helps in some respects to
bridge the gap between the two groups. Taylor’s (1918) figure of a hypopygium
referred here by Edwards (1924) was actually that of A. wilsoni Tayl. Described
from twelve specimens, all from EHidsvold.

Allotype ¢: Hidsvold, S. Queensland, Oct. 13, 1926 (Mackerras), in the Macleay
Museum, University of Sydney.

This species was abundant at Hidsvold from July to October, 1926, though
usually a rarity in that district. It is a day and night biting sylvan species,
occasionally also taken in the scrubs. The early stages were not discovered.

Distribution.—‘An inland species . . . . widely spread from Queensland
to Victoria and into South Australia’ (Ferguson, 1926). Though in general
distributed west of the Divide, it appears in the coastal districts in south
Queensland.

AEDES (OCHLEROTATUS) THEOBALDI (Taylor) form EIDSVOLDENSIS, new form.

®. More ornate than the typical form from which it differs in the following
respects: the pale scales of the wings, though as numerous, are distinctly narrower;

296 NOTES ON AUSTRALIAN MOSQUITOES, ii,

the thorax is densely covered with small, dull creamy golden scales without darker
admixture; abdominal tergites with broad transverse basal creamy bands produced
posteriorly to form a prominent median triangle which increases in size on the
posterior segments, until on segment six it forms a complete vitta; the mottling
of the darker parts of the segments is hardly detectable. This is possibly a distinct
species, but I hesitate to treat it as such in view of its closeness to A. theobaldi
Tayl. and the fact that only one specimen was available for study.

Holotype: The unique female, taken biting by day in forest country at Hidsvold,
Oct. 7, 1926 (Mackerras), has been lodged in the Macleay Museum, University of
Sydney.

AEDES (OCHLEROTATUS) NORMANENSIS (Taylor).

This obscure and difficult species is very close to A. theobaldi Tayl., but is
even more likely to be confused with A. vigilax Sk. from which it is abundantly
different on hypopygial characters. It is distinctly smaller than either and may
be separated from both by a collective consideration of the following characters :—

1. The pale scales of the wings are narrow and are only numerous along the
anterior border. On the rest of the wing they are very scanty or absent, though
a few are almost always present along Cu.

2. The scutum bears relatively large pale golden scales without darker
admixture; they are not densely packed and show a tendency to a linear
arrangement.

3. The basal pale bands of the abdominal tergites are variable in extent,
being occasionally conspicuous and straight edged, but more often reduced to a
narrow patch. There is no mottling.

4. There is a variable amount of irregular pale scaling on the proboscis
beneath. It shows no sharp demarkation from the distal dark part.

5. The mottling of the femora is relatively inconspicuous and the last hind
tarsal segment is devoid of, or with only a narrow, pale ring.

The characters of the male are even less definite than those of the female
except for the hypopygium. This structure (Text-fig. 6) shows close affinities with
that of A. theobaldi Tayl., from which it differs in the smaller but more definite
apical lobe and the differently shaped basal lobe of the side-piece.

A. normanensis is a day biting forest species which is numerous at Hidsvold
in the later half of summer but is rare earlier in the season. I have bred it from
mixed pupae taken in a muddy rock pool and in a waterhole in the bed of a
sandy creek. The larvae are extraordinarily elusive and could not be found.

Distribution.—South and Central Queensland.

AEDES (OCHLEROTATUS) VIGILAX (Skuse).

A re-examination of Skuse’s type leaves no doubt as to the identity of this
very abundant and annoying insect. It is also clear that it was correctly
recognized by Edwards, despite the fact that he states that the wings are entirely
dark scaled. The following notes made from the type will serve to separate
A. vigilax Sk. from any of its allies; wings with scattered pale scales of normal
form, fairly numerous along the anterior border, some scattered ones on the
other veins, particularly along Cu; scutum with fine scales, mostly dark brown,
but interspersed with numerous irregularly arranged bronzy golden ones; pale
bands of abdominal tergites white, about one-fourth the width of the segments
and straight edged; pale area on ventral side of proboscis conspicuous and with

BY I. M. MACKERRAS. 297

sharply defined apical limit; femora mottled with numerous pale scales anteriorly;
last hind tarsal segment with a broad prominent basal white ring; lower
mesepimeral bristle absent. In over 100 females from various localities the range
of variation is slight, being practically limited to the amount of pale scaling on
the wing; the white scales may be completely limited to the anterior border, or may
be so numerous as to approach the condition seen in A. theobaldi Tayl. The
apical lobe of the side-piece of the male (Text-fig. 7) is not apparent and the basal
lobe is large, rounded and covered with numerous hairs, the pigmented bases of
which give it a dark appearance; the claspette is small and bears a simple straight
apical bristle representing the appendage.

The larvae have been described by Cooling (1924) and Brug (1924) and the
larva and pupa figured by Hill (1925a). Their most characteristic features are
the short broad siphon, the arrangement of the comb scales in a patch, the short
antennae, and the short frontal hairs.

A. vigilax Sk. is the most important of the Australian “bush” iosqniteck:
It is exceedingly abundant in the vicinity of estuaries where there are any mud
flats or mangrove swamps which appear to be its principal breeding grounds and
the dominant factor governing its local prevalence. In such localities as the
Hawkesbury River it may settle on one in clouds and make life unbearable. It
has, however, a wide range of flight and may be quite numerous several miles
from its breeding grounds. It bites both by day and by night. Its habits and the
question of its control in the Sydney district have been dealt with elsewhere
(Mackerras, 1926); they do not appear to differ materially in other parts of its
range.

Distribution.—Widespread round the coast of Australia. Though almost
exclusively coastal in distribution, I have taken a few specimens well away from
salt water, one at Barrington Tops, N.S.W. (January, 1925) at 5,000 feet elevation
and 40 miles from the coast, and several at Eidsvold, S.Q. (December, 1922,
April, 1924, and October, 1926). It appears certain that it was breeding in the
latter locality if not in both. The season is from October to April, the greatest
prevalence being in the middle of summer.

AEDES (OCHLEROTATUS) RUBRITHORAX (Macquart).

A relatively rare species which is easily recognized by the entire absence of
pale scales on the anterior aspect of the femora and on the wings, and by the
reddish scutum which is covered with fine bright golden scales. The male
hypopygium (Text-fig. 8) resembles that of A. vigilaz Sk., differing only in the
broader, more rounded side-piece bearing a more prominent basal lobe which is
not darker in colour than the rest of the side-piece.

Edwards (1924) queried Culex procax Skuse as a synonym, an opinion which
was confirmed by Ferguson (1926) who also pointed out that one or two weak
lower mesepimeral bristles may be present. These findings are confirmed by a
re-examination of Skuse’s type and the study of a fair series of the species
commonly known as A. rubrithorax Macq. The bristles may be present on one
side and absent on the other; when present they are small but quite conspicuous.
The fact that Culex rubrithorax Macq. was described from Tasmania need not
affect the determination, since it is now known that many species described as
from Tasmania in Macquart’s 4th Supplement actually came from coastal New
South Wales: His description fits an abraded specimen reasonably well and it
seems preferable to use his name rather than Skuse’s, although the latter can be

298 NOTES ON AUSTRALIAN MOSQUITOES, ii,

definitely recognized and the type is in good condition. In the same work
Edwards also queries Culex occidentalis Skuse as a synonym. The type of this
species, is, however, a Finlaya and is in my opinion identical with A. (F.)
queenslandis Strick.

Distribution.—A coastal species extending from South Queensland to Sydney,
New South Wales; also recorded, almost certainly erroneously, from Western
Australia and Tasmania. Specimens are before me from the following localities:
Hidsvold, 8.Q. (February and April); Stradbroke Is., S.Q. (September); Myall
Lakes, N.S.W. (August); North Harbour, Sydney (October); Balmoral, Sydney
(November). Purely a bush species and a day biter.

References.

Brug, S. L., 1924.—Notes on the Dutch-Hast-Indian Mosquitoes, Bull. Ent. Res., xiv, 4,
pp. 433-442.

Cootine, L. E., 1913.—Report on Mosquito work in Brisbane, Ann. Rpt. Commissr. Pub.
Hlth. Qid., to 30th June, 1913. Appendix J, pp. 55-64.

,1924.—The larval stages and biology of the commoner species of Australian
Mosquitoes, with the biology of Aedes pecuniosus Edwards. C’wlth. Aus. HIth.
Dept. Serv. Publn. (Trop. Divn.), No. 8, pp. 1-40.

Epwarps, F. W., 1912.—A key to the Australian species of Ochlerotatus (Culicidae).
Ann. Mag. Nat. Hist., ser. 8, ix, pp. 521-537.

, 1922a.—A synopsis of the adult Oriental Culicine (including Megarhinine and
Sabethine) Mosquitoes. Part II. Ind. Jl. Med. Res., x, 2, pp. 430-475.

, 1922b.—A revision of the Mosquitoes of the Palaearctic Region. Bull. Ent.
Res., xii, 3, pp. 263-351.

, 1924.—A synopsis of the adult Mosquitoes of the Australasian Region. BuwIll.
Ent. Res., xiv, 4, pp. 351-401.

, 1926.—Mosquito Notes.—VI. Bull. Ent. Res. xvii, 2, pp. 101-131.

FpRGUSON, E. W., 1926.—Mosquitoes of New South Wales. 15th Rpt. Microbiol. Lab.,
N.S.W., for 1924, pp. 187-191.

HarRRISON, L., 1927.—The Composition and Origins of the Australian Fauna, with special
reference to the Wegener Hypothesis. Presidential Address to Section D, Aust.
Assn. Adv. Sci., Perth, 1926 (in press).

Hiuu, G. F., 1922.—Notes on the Habits and Distribution of some North Australian
Culicidae. C’wlth. Aus. Hlth. Dept. Serv. Publn. No. 21, pp. 1-40.

, 1925a.—The distribution of Anopheline Mosquitoes in the Australian Region,
with notes on some Culicine species. Proc. Roy. Soc. Vict., n.s. xxxvii, pp. 61-77.

, 1925b.—Mosquitoes of Australia. Vict. Natwralist, xlii, 2, pp. 45-46.

MacKerRrRAS, I. M., 1926.—The Mosquitoes of the Sydney District. Presidential Address.
Aust. Naturalist, vi, 3, pp. 33-42.

Taytor, F. H., 1918.—Contributions to a knowledge of Australian Culicidae. No. iv.
Proc. Linn. Soc. N.S.W., xliii, pp. 826-843.

NOTES ON AUSTRALIAN DIPTERA.* No. xi.

By J. R. MALLocH. NWS i
(Communicated by Dr. I. M. Mackerras.) ae

(Twenty-three Text-figures. )
[Read 27th July, 1927.]

Family Calliphoridae.

As an inducement to the study of this family by Australian workers I offer
some preliminary notes on the family characters and groupings within the family,
and also present descriptions of some new Australian genera and species derived
from material submitted to me for identification by the late Dr. Hustace W.
Ferguson. Many of the species are of considerable economic importance, and
though most of them are widely distributed and common, definite specific
identifications are difficult to arrive at by the use of existing descriptions and the
few keys available. Later on, should material become available to me, I may be
able to present a fuller revision of the Australian species.

The first difficulty that confronts the systematist in this, as in all other
taxonomic work, is the definition of the group or family. To state this may appear
superfluous, but the more one widens his acquaintance with taxonomic treatises
the more surely he is forced to the conclusion that many, if not most, authors
possess merely an intuitive knowledge of the limits of the group they write of, or
their concept when reduced to plain English is so broad, ambiguous, or so full of
exceptions, that the uninitiated student is largely or completely in the dark as to
what really constitutes the group. This is largely true of the status of the work
on the family Calliphoridae to-day. Some workers include in it the Sarcophaginae,
while others exclude that subfamily, consider it a family, and incline to break the
residue up into families or subfamilies according to individual taste or opinion.
Personally I prefer to consider all genera that have the second abdominal sternite
overlapping sides of the tergite, no well developed convexly formed postscutellum,
which possess strong hypopleural bristles in one or more vertical series below the
spiracle, and have the first posterior cell of the wing conspicuously narrowed at
apex as belonging to the family. This definition would include Calliphorinae,
Sarcophaginae, Rhiniinae, Chrysomyiinae, Metopiinae, and some other groups
which may be of equivalent or lesser rank such as are represented by
Mesembrinella, Bengalia, etc., but none of which occur in Australia so far as I
am aware at this time.

1[Mr. Malloch has given the references to Macquart’s papers on Exotic Diptera to
the original published in ‘“‘Mémoires de la Société des Sciences, de l’Agriculture et des
Arts de Lille’. As these Memoirs are not in Sydney, I have added the references to the
reprint published in Paris which is in the library of the Society. For a list of dates of
publication see Cat. Library Brit. Mus. iii, L-O, p. 1215, 1910.—EHp.]
M

300 NOTES ON AUSTRALIAN DIPTERA, Xi,

At present I possess Australian representatives of all of the five larger groups
listed above, and below I give a key for the recognition of these in the hope that
it may prove useful to students of the calyptrate flies. It is to be noted, however,
that this key is intended to apply to Australian forms only. Some extralimital
forms, such as the genus Compsomyia, which does not have erect hairs on the
small knob below the base of wing, will not run out to their proper places in
the Key.

Key to Subfamilies of Australian Calliphoridae.

1. Stem-vein of wing (radius) not setulose on posterior upper side of its basal section 2
Stem-vein of wing with distant setulae or hairs on posterior upper side of its basal

SOCELOME race GCOS are eH aire) kn cly eae os alee RSA SIO CARS RST A PREC price ht us) 1A TEN 2 ca 5
Lower calypter with some long fine erect hairs on a part of its disc above ..........

BERN REA Cr SL ERAS pees ht Store, Ohh rT ROSE EREORCEETE CRE CNS Geola sO GOO bad ona Calliphorinae, pt.
Lower calypter without long erect hairs on any part of its disc above .......... 3
8. Arista bare or very short pubescent; notopleural bristles two in number ..........

to

ACER CN TERE ORES GN O NC cies CAE NOT RPS SL FeROR HENAN O ESAIOl cee nORe nak ras eC RCmIC Cumiey Gb G Gi co" ot ORI RANGA ReneeiS Metopiinae
Arista with at least short hairs, the longest distinctly longer than width of arista
BENDS Sis Eo he vesisyiey arlene ewer oka relies Mush ekaucole cise Re esl ee elise we coals chattc uate ce oR RMeSNR e eat ec) so Se 4

4. Notopleurals nearly always two in number; the outer two bristles on area between
the presutural dorsocentrals and notopleurals not in a straight line, the anterior
one distinctly laterad of the posterior .....................- Calliphorinae, pt.

Notopleurals almost invariably three or four in number; the two outer bristles on
the area between presutural dorsocentrals and notopleurals either in longitudinal
line, or anterior one mesiad of posterior .............2c0eeeeeees Sarcophayginae

5. The small rounded protuberance below base of wing frequently with erect hairs, lower
calypter subtruncate at apex, concave on outer margin, haired on part of upper
SUPEA GCOS chy eites te calicestens tairettove ite cee ia te weedemotlairer inn each oiramone wreus, ssi tehepenc iM onee Mem Rom CHS Chrysomyiinae

The small rounded protuberance below base of wing, and upper surface of lower
calypter, bare, or without distinct hairs, the calypter rather narrowly rounded
at ‘apex, straight; or nearly so; on) outeriomargin joanne oer acticin. Rhiniinae

In connection with the above key it appears pertinent to point out that there
are some characters in the structure of the male hypopygia which may have a
group significance. I do not incline to give much weight to the structure of these
organs in distinguishing genera or higher groups as a general rule but in some
families the general structure may sometimes be relied upon to identify the males
belonging to particular genera. If a comparison is made between the hypopygium
of a species of Calliphora and one of Sarcophaga it is evident at a glance that there
is a very decided difference in the two pairs of claspers which are directed
downward at apex. In Sarcophaga the outer pair are short and very much less
conspicuous than the inner pair, being usually referred to as the accessory plates,
while in Calliphora they are usually as long as the inner pair and quite as heavily
chitinized. There are some other distinctions but there is so much variation in
the inner portions involved in the penis and its guard in the genus Sarcophaga
and its allies that little use can be made of these in generalizing.

Undoubtedly competent morphologists could homologize the parts of the
hypopygia but the taxonomist is primarily interested in differentiations and not in
homologies so occasionally the latter group uses different names for the same part
of an organ which course, while probably perfectly justifiable from the point of
view of one more especially interested in obtaining identifications than in arriving
at homologies, leads to some confusion in terminology.

I have figured the hypopygia of some of the other genera dealt with in this
paper and a comparison of these with the figures presented in some of the papers
on Australian Sarcophagidae will readily show the distinction to which I refer.

BY J. R. MALLOCH. 301

One of the fatal objections to the use of the structure of the male hypopygia
as an index to generic and subfamily relationships is that it is very rarely the
case that co-ordinated characters can be found in the females and a classification
based upon one sex is, not only a hindrance, but practically a bar to the study of
any group in which it is so used unless one has authentic males and females of
the species, which it is rarely possible to obtain without long and arduous
field work.

Subfamily Calliphorinae.

In this subfamily there are in use several generic names which have rather a
doubtful status, the basis for the separations being the different number of the
so-called sublateral bristles present. These consist of from one to three bristles
in front of the thoracic suture in line with the postsutural intra-alar bristles and
mesiad of the two presutural bristles referred to in the foregoing key to genera, the
latter being known as the posthumeral (anterior) and presutural (posterior) bristles
respectively. The genus Cynomyia, which I have not seen from Australia, has but
one of these sublateral bristles, which is almost in line transversely with the hind
margin of the humeral callosity, both the weak anterior one and the one just in
front of suture being absent. In the genus Steringomyia, also unknown to me
from Australia, there are two of these bristles present, while in Onesia and
Calliphora all three are usually present.

If we apply these characters to the Australian species included in Calliphora
by Patton and by Hardy in their recent papers it becomes evident that there are
either two or more genera present, or the character is not of generic value.
Cynomyia I consider is a good genus, but in some Australian species of Calliphora
the posterior one of the sublateral bristles may be entirely indistinguishable in
some specimens, while it is evident though weak in others even of the same
species. I am inclined to consider that under these circumstances the Australian
species at least cannot be thus generically divided. I find that in Onesia there
are from one to three sublaterals present, the usual number being three. I do not
care to express an opinion here on the validity of Steringomyia.

The distinctions between Onesia and Calliphora are even less marked than
are those between the latter and Steringomyia. The most recent attempt to
separate these is that by Shannon in 1923 in dealing with the North American
forms. He cites the straight or slightly curved apical section of fourth vein as
against the distinctly curved one, and the short as against the longer third
antennal segment as characters in Onesia and Calliphora respectively, but there
is no reason to accept these as generic criteria, the evident intent being merely to
perpetuate, and not to discard existing generic concepts. The worst feature of
accepting Onesia as the same as Calliphora is that the former has page priority
in the paper in which they both were published and thus would supplant
Calliphora, with a consequent change of the name of the family to Onesiidae.

It is unfortunate that intensive research almost invariably results in more
or less radical changes in names of even the most widely known and common
insects, and the fault lies not with the modern worker who applies the rule of
priority, but with those of past generations who failed to appreciate identities
and co-ordinate related forms. Were it possible to accept Onesia and Calliphora as
distinct I would not hesitate to suggest this course, but I can see no good excuse
for doing so on any characters distinguishable by me.

En passant it must be remembered that we have been using the name Calliphora
contrary to the rules of priority, as it has been proven that the name properly

302 NOTES ON AUSTRALIAN DIPTERA, Xi,

applied to the group is Musca, and that the group now known under the generic
name Musca should be known as Promusca Townsend. An attempt has been made
to conserve the name Musca for the group containing domestica and its allies,
and in the event that such a course is adopted then the name Calliphora will
naturally fall as a synonym of Onesia. The proper medium for the final disposition
of these points would be a monograph on the family so I merely state conditions,
and do not make any drastic changes herein, this paper being intended merely to
stimulate interest in the family it deals with.

I present below a key for the separation of the few genera of this subfamily
I have seen from Australia. There are probably more native genera, but the
amount of material available to me now is rather scanty, and possibly with this
paper in hand some student of the calyptrate flies may be induced to take an
interest in it so that more specimens from a wider range of territory may become
available.

Before I attempt to deal with the genera and species of this family it may be
well to give space to some remarks upon the status of some of the names in the
group apart from the generic names just referred to.

Major Patton has been for some years actively engaged in publishing papers
on Calliphoridae and Muscidae, particularly those forms which affect man and
domestic animals, and one of his recent papers deals with Australian Calliphoridae.
In this paper he cites a number of synonyms and describes several new species.
All of the new species are available to me in material sent to me by Dr. Ferguson,
so that it is possible to include data upon them in the present paper. It is, however,
impossible for me to decide definitely whether his conclusions as to specific
identities are or are not correct. I incline to the opinion that some at least of the
species he includes as synonyms are perfectly valid species, but without having
the same opportunity as he had to examine the type specimens it would be
unwise to be dogmatic in giving determinations upon most of these. Mr. Hardy
has also recently published a paper dealing with the species of Calliphora and
though his ideas and mine appear to conform better than do Major Patton’s and
mine there are so many additional species in my material which he did not have
that it is impossible for me to definitely decide what certain of his species are
in my lot. However, it may be possible after my paper appears for the data
presented to be used in bringing together material for use in a comprehensive
study of all the Australian species of the family.

The most recent paper dealing with Calliphoridae is that by Mr. Senior-White.
This presents a consideration of the Oriental species, but unfortunately has much
the same blemishes present in Patton’s work, which is to be expected because
there was a close collaboration between these workers. I note here that Calliphora
fulviceps van der Wulp, an endemic Oriental species, is omitted by Senior-White.
This is a very characteristic species, with bright fulvous yellow face and cheeks,
the latter furnished with long fulvous hairs, and it has but one pair of presutural
acrostichals as a rule, a character which I have not seen elsewhere in the genus.
I have not seen this species from other localities than the mountains of Java
and Sumatra.

Key to Genera of Calliphorinae.
1. Lower calypter with some conspicuous erect fine hairs on at least part of its upper
suriace, sometimes yery close to ibase,. omy, ra ctopschotensie:ch orci «lolosiis’sloretc cakeborene 2
Lower calypter without any erect fine hairs on upper surface .............esce0eee 4
2. Extreme base of stem-vein of wing on under surface with setulose hairs; upper
surface of lower calypter almost entirely haired; presutural area with 3 to 5
bristles

BY J. R. MALLOCH. 303

Extreme base of stem-vein of wing without distinct hairs or setulae; apical part of
upper surface of lower calypter broadly bare .... Calliphora Robineau-Desvoidy

3. Byes hairy; no strong forwardly directed orbitals in female; the fourth tergite in
this sex with a central longitudinal weak part which gives the segment the

SHOVE WCENAEE) GME |axeTbNS USNS GoooohobuaocgoadasnouogdDodGaUcKUorK Ptilonesia Bezzi.

Eyes bare; two strong forwardly directed orbitals on each side in female; fourth
tergite in this sex entire, the apex transverse and heavily chitinized ............

Mee Te rape pray ciaaN Puma lotauieseneces aides sve tevers user cuecaraue (auesssieCalivceue ¢cs eilecenesa Xenocalliphora Malloch

4, Suprasquamal ridge strongly setulose anteriorly and posteriorly ..............+502008
RRSP eat WN Merce ele bay ty hecq atti VAP inci lek iets e Al s M ly Lucilia Robineau-Desvoidy
Suprasquamal ridge bare at least anteriorly or posteriorly .................... 5

5. Supraspiracular convexity of metathorax with long erect fine hairs ................
A eE a AUR EHO RECS Sone GeO) Gy ADIL BUS SCOCSSU DESC EERPaISTEHERE. Hise Pee ne weaae tie aE Paratricyclea Villeneuve

Supraspiracular convexity without erect fine hairs ..................0 00s ences 6

6. Propleura haired in centre between the humeral hairs and those on lower margin .. 7
ProOpleura ware Oy CNEE Cae atlas ei acwlio Susu alens sicucuedel miaieee lena uaiepeneusheral seis tehjols anens isvelsieiacensers 9

7. Presutural area with 4 or 5 bristles ............... Melinda Robineau-Desvoidy, pt.
Presutunalvarea withitw Ov GiStlesivarccce ctovcie ce eral cieccneyeneie ele lore sis) oleieiteriens lollelisiialelteele cls tel iaele 8

8. Upper half of parafacials with setulose hairs; supraspiracular ridge bare at posterior
OURS TINT Yea eNe, cle cass laerte late aies eee or eney Sirona aitetisuetiode ohalieneuts Melinda Robineau-Desvoidy, pt.

Upper half of parafacials bare; supraspiracular ridge fine haired at posterior extremity
re Nene reece ee toa ras alten station ace cs caungalea eel tahes eremacenetanre ameiotanena lene tacartee omepe tones Euphumosia Malloch
9. Thorax without yellow or golden pile or tomentum besides the usual bristles and

PANT SUP a Tepe OO spac iag shia pen ea rotten neue doy ate naka pollens) oF ah snaila’ succoynsagey esueaebe tate Anthracomyia, gen. n.
Thorax with yellow or golden pile or tomentum in addition to the usual bristles and
VATS aie ecauersne be oh Meta ae deme en SUA rte SinWek su seem rer urey are uecaiene accra rele Pollenia Robineau-Desvoidy

Genus PriLoNEsIA Bezzi (1927).
Bull. Ent. Res., 1927, xvii, p. 242.

This genus contains one known species, with characters as indicated in the
preceding key. It is most closely related to Xenocalliphora Malloch, an austral
genus known from New Zealand. I prefer to treat this as a genus and not a
subgenus as Bezzi has done. Genotype, Pollenia auronotata Macquart.

PTILONESIA AURONOTATA (Macquart).

Pollenia auronotata Macq., Mém. Soc. Sci. Lille, 1854, p. 135 (1855); Dipt.
Exot., Suppl., 5, p. 115, 1855.

Similar in general habitus and colouration to Calliphora erythrocephala, but
the bright orange coverings of the thoracic spiracles, similarly coloured scales
over wing base, and the entirely different armature of the head readily distinguishes
the species. The calyptrae are fuscous, with white edges.

Length, 9-10 mm.

Locality: Sydney, N.S.W., one female.

Genus CaLitipHorA Robineau-Desvoidy (1830).

The Australian members of this genus present a greater diversity of form,
colour and vestiture than do those known to me from any other faunal region,
and it is not surprising that certain authors have selected some of the species as
types of distinct genera. In the case of genera acceptance or rejection is more a
matter of individual opinion than is the acceptance or rejection of species.
Almost invariably it is possible to demonstrate the existence or absence of
differentiating characters in species, but in accepting or rejecting genera one has
to decide whether the possession of one or more distinguishing characters in
any species or group of species is sufficient grounds for considering them as
entitled to generic separation from their most closely allied relatives. Some
authors incline to accept a single structural character, such as the presence or

— 804 NOTES ON AUSTRALIAN DIPTERA, Xi,

absence of hairs on the eyes, as sufficient to warrant the erection of a genus for
the reception of species previously included in a genus the species of which do
not possess that character, even when the whole of the species are otherwise
similar in features distinguishing them from related genera. The trend of this
class of work is towards the erection of a great number of monobasic genera

Acrostic
Pa | y aan
' Re | ii
f P poy \ q
stil data
Sn
i

Cas
—

Pastsvtvural |Resutura

Text-figure 1. Head from the side, typical. An, antenna; Ar, arista;
C, cheek; HE, eye; F, face; Fr, facial ridge; In, interfrontalia; Inf, inferior
orbital bristles; Oc, ocelli; Ocb, ocellar bristles; Or, orbit; P, palpi; Pf,
parafacial; Poc, postocular cilia; Pr, proboscis; Pv, postvertical bristles; Sup,
supraorbital bristles; Vb, vertical bristles; Vi, vibrissae.

Text-figure 2. Head from the front, female. Lettering as Text-figure 1, with
additions, Af, antennal foveae; Ft, frontal triangle.

Text-figure 3. Dorsum of thorax, bristles lettered.

Text-figure 4. Side of thorax, no definite genus. Be, basicosta; Bs, basal
sclerite of stem vein; C, coxae; H, hypopleura; Hu, humerus; K, knob-like
pleural process; Le, lower calypter; M, mesopleura; P, pteropleura; Pb,
propleural bristle; Pr, propleura; Sb, stigmatal bristle; St, sternopleura; Suc,
supraspiracular convexity.

without materially assisting the student interested in classification. In other
words, monobasic genera tend to obscure relationships, a result that is less
desirable than is the retention in one genus of species biologically similar but
separable by such a character as that mentioned above. I do not think that the
genera Neopollenia Brauer, Neocalliphora Brauer and SBergenstamm, and

BY J. R. MALLOCH. 305

Paracalliphora Townsend, are entitled to full generic rank, and include them, or at
least their genotypes, in my synopsis given below.

I have found it impossible to determine definitely whether or not certain
forms in my possession are distinct species without a dissection of the male
hypopygia and present drawings of these herein to facilitate recognition of the
species involved by other students of the group. These organs are not so intricate
as those of most species of Sarcophaga, but the interior central portions, consti-
tuting the penis and its sheath, etc., differ more or less markedly in the different
species, providing characters by means of which they may be recognized. It is
well known that the male hypopygia are of great value as specific criteria in many
groups of insects and in the Diptera such families as Sarcophagidae, Chironomidae,
Tipulidae, Dolichopodidae, Asilidae, and others are striking cases in which these
organs provide reliable and outstanding means of identification in the male sex.
It is not so well known, however, that some families and genera do not present
great differences in the structure of the hypopygia of the males, and one need only
-Mmake a study of the males of the genus Musca sens. lat. to discover how trivial are
the structural differences between the hypopygia of even the most distinct species
of the genus. In fact, Major Patton has expressed the opinion that these organs
offer no means for distinguishing the species. This is not strictly correct, but the
distinctions are so trivial that only one versed in the differentiation of species
can appreciate them. In Calliphora the structure of the two apical pairs of claspers
or forceps is very much the same in many of the species, but there are minor
differences of form and armature, which, coupled with the differences in the
structure of the central portions, readily distinguish the species I have examined.

To facilitate the use of this paper I give diagrams showing the portions of
the thorax and the position of the various bristles utilized in classification
(Text-figs. 1-4).

Key to the Species.

1. Eyes in both sexes with quite dense erect yellow hairs ...................2008- 2
yes! in’ neither ‘sex with: noticeable hairs) see jsc sce sce ees reece cern cals 3

2. Thoracic dorsum densely yellowish dusted so that the ground colour is entirely
or almost entirely obscured and the thorax appears entirely ochreous ......

RPV APED LR LIAE REL IS 0 Ee ChB. ue wets PACER ror een Ie ieee nan OCILTCLCe Gm SCHINET
Thoracic dorsum bluish black, with whitish dusting which does not at ail obscure

lO). Fseroyonael oVloybhe” GooonedodcosccsdcbcuuLoccuS ochracea form nigrithorax nov.

3. At least the tibiae reddish or yellowish, and distinctly paler than femora, or the
TOMO, ES BYISo APO? cooaoccodoob oop booU Rad OOOmOD DO DONO ODDO ODO aDD0OS 4

Legs entirely black, tibiae rarely very slightly paler than the black femora ...... 8

4. Pleural and abdominal hairs and bristles entirely black; abdomen black, with
bright golden yellow pollen which is noticeably checkered; femora, tarsi, and
coxae, black, tibiae reddish, darker at bases; posterior sublateral bristle
VVC Rar Hh SUSAR A TT STS RROD ET, en hearers Ae rah hace holed a ihren eka ate areronte tibialis Macquart

Pleural and abdominal hairs in part bright yellow or golden .................... 5

5. Abdomen black or fuscous, with a metallic bronzy tinge and dense yellow dusting
which is conspicuously checkered as in tibialis, the ventral hairs longer, and,
like the lateral hairs and bristles, bright golden yellow; posterior sublateral

DEIstlewusuallystGistinets 44a ees cre sete eno atone eh sre ohne che nto een tale Gictal cloth aie cece reteset dA
Abdomen honey yellow on at least the sides and venter, always with a large
part of central section of disc metallic blue, or violet coloured .............. 6

5A. Eyes of male with facets of upper half conspicuously larger than those of lower
half, the frons reduced to a narrow line; thorax almost invariably with three
pairs of presutural acrostichals; hypopygium as Text-figure 5 ............
OF AOTO UDO CIELO OGRE RCAC RELDRCR CRB ICROR LTE SICH EUS ESIO STE NOLS LER SOIC RRAC IO RG Co eae stygia (Fabricius)

306 NOTES ON AUSTRALIAN DIPTERA, Xi,

Byes of male with facets of upper half but little larger than those of lower half,
the frons about as wide as third antennal segment; thorax almost invariably
with two pairs of presutural acrostichals; hypopygium as Text-figure 6 ....
SS ETI en ISL Hitt ATES AER Gleick ORE Die TOC ORO HIG TEN GRY cl eck Cth OPER EMCEE ERECT Cron hil Patton

6. Coxae, femora, and bases of tibiae, dark brown or fuscous; thoracic dorsum when
seen from behind with a narrow postsutural vitta between the acrostichals,
and a spot between these and the dorsocentrals midway between suture and
posterior margin, black; scutellum with the pruinosity distinctly checkered,
best seen by moving the specimen round in examination; dorsum. of abdomen
with most of disc of basal three tergites glossy purplish or violet coloured, the
dark portion not sharply limited, fourth tergite densely golden dusted, slightly

checkered; posterior sublateral bristle lacking ............ fuscofemorata, nN. sp.
Coxae, femora, and tibiae, bright honey yellow; dorsum of thorax and scutellum
IGE WAAL GIS AWE oooccdovoonnuocoooadgogoucccodcoousoooudcouDObOOUOS 6A

6a. Thorax with two pairs of presutural acrostichal bristles, the posterior pair
distinctly proximad of a transverse line drawn between the posterior pair of
presutural dorsocentrals; posterior sublateral bristle lacking or almost indis-
tinguishable from the surrounding hairs; fourth abdominal tergite densely

yellow dusted, with a large underlying metallic greenish basal mark which is
obscured. by tHe dusting 1) eee is ec ase e sitetiovot ee oehate reas augur (Fabricius)

Thorax with at least three pairs of presutural acrostichal bristles, the posterior pair
just in front of suture, and much behind posterior presutural dorsocentrals;
posterior sublateral bristle present, quite distinct from the surrounding hairs 7

7. Humeri bluish black, concolorous with dorsum of thorax; dorsum of abdomen
broadly purple on all tergites, the dark colour of fourth conspicuous because of
the presence of but thin whitish dusting; sternopleurals 1+1... macleayi, n. sp.

Humeri testaceous, very noticeably paler than dorsum of thorax; dorsum of abdomen
partly metallic green or blue-green; fourth tergite with more conspicuous

dustinge’; jsternopleurals %2) 70h) suis jaeetedarec eterstel eeiaicnemevel el or eiicpeuenevouers centralis, n. sp.
8) ‘Bosterion sublateralgbristlema cine cwvsii sires iene EMI eater ie reiee nets 9
Posterior sublateral bristle present, sometimes weak and short .............. 10

9. Small species, about 5 mm. in length; abdomen bright metallic blue-green, almost
entirely lacking whitish dusting on dorsum; wing veins entirely fuscous; apical
section of fourth vein beyond the angle almost or quite straight ........ sp. ?

Larger species, about 10 mm. in length; abdomen brassy green, with rather dense
whitish dusting which is distinctly checkered; wing veins yellow at bases, brown
beyond; apical section of fourth vein beyond bend distinctly curved ........
OOOO RR OIC LCE EL RRR IE Teaco ITO nora Glo. od yo Ea atolo dG dispar Macquart?

10. The small unarmed scale (basicosta) between tegula and base of costal vein black

OVE NER SCOUS) bys 5545 ay sho chsiaiecices sakes ee ad tod See Bana Saas) Sch HOD ote Levi ae aOR eet pray athe tratve re Betoionter le matse events, ouielle 11
The small unarmed scale between tegula and base of costal vein whitish or yellowish
at) least ron outer halfuior AM One: naan Wobvals sae enc ie varalati mene Rove Ponte retekaterlotieirodekare varteiralielte Lome 14

11. Abdomen dark metallic blue, with slight even dusting and a dark central vitta
on basal three visible tergites, the fourth with quite dense brassy or yellowish
dust, sharply contrasting with the other tergites; calyptrae dark brown or
fuscous, fringe of lower one yellowish; hairs on occiput except postocular fringe

yellow; those: on cheeks iDlacky si eieysslousde cheteeees eee tees s teens apicalis, n. sp.
Abdomen metallic blue-green or black-blue, with or almost without whitish dusting,
fourth tergite not at all differentiated from the others .................... 12

12. Robust species, with dull metallic blackish blue abdomen which is quite noticeably
white dusted; scutellum with 18-20 bristles; hypopygium as Text-figure 7

SRiend Coe eo an Game boom One OOS ODODE CUD DU oD b OO fone d 06.80 a DOO r OO robusta, n. sp,

More slender species, with brilliant metallic blue-green abdomen which is but
slishtly white dusted’; scutellum with) 10-12) sbristles) Wiis. yas. cas sci eeiene 13

13, Outer forceps of male hypopygium stout (Text-figure 8); third antennal segment
orange, slightly darkened ‘at apex! .jds).sw scenic tiein es clalenetone «2 xanthocera, n. sp.

Outer forceps of male hypopygium slender and tapered apically (Text-figure 9) ;
third antennal segment fuscous, narrowly yellowish at base .. australica, n. sp.

14. Lower calypter fuscous, with a portion of margin as well as the thickened edge and

the fringe, white; cheeks and lower portion of parafacial brownish orange, with
black hairs; fourth vein beyond apical curve much bent .. erythrocephala Meigen

15.

16.

17.

18.

184A.

18B.

AL),

20.

21.

22.

BY J. R. MALLOCH. 307

Lower calypter brownish, yellowish, or whitish, if the margin is paler than the
disc the pale colour is confined to the thickened edge, and does not extend over
any part of the disc; cheeks with the raised portion black or fuscous, never

orange, if rufous the surface is obscured by grey dust ................-.. 15
TW ENLACE © Pasi ote sho Gb 10. cic: COLE. CRORN COT oTe rr nGrG o OiaiG b Slots Bidibro GucKo oro +O ORT ONO LDF OLD tO: 0 ia OO, CIC IOLEVO IO 16
TM esGaN KS) Wy leo ote Sous Gla COI CIO EIA ODEeTO'G Cho GIG OOD, CuOTE CLCRO CIOrG & DDANO ORO CS ORO OC Onn OO ODIO OFGIO TO 19

Facets of upper half of eye larger than usual, the frons reduced above to a very
narrow line, which is not wider than anterior ocellus; dusting on entire orbits
and parafacials uniformly yellowish grey and dense, when seen from the side
without a dark patch opposite base of antennae and another below middle of

parafacial; hypopygium as Text-figure 11 ...................-.. plebeia, n. sp.
Facets of upper half of eye not especially enlarged, the frons at narrowest point at
leastnuwicesasi wider as anterlormoCellUsiserercuce cisicusiellelelelel en -iclienehalielloceielel oSellalionsis 17

Frontal orbits and parafacials densely and uniformly yellowish grey dusted, not
noticeably checkered when seen from the side; apical section of fourth vein
almost straight beyond angle; hypopygium as Text-figure 10 .... minor, n. sp.

Frontal orbits and parafacials variably dusted, with a noticeable white patch on
latter just below bases of antennae which is best seen when the head is viewed
from the side; if orbits are evenly dusted, the apical section of fourth vein is
DS TUERPY RARE re evar itins RE SIA CRO SS RAOLR 5. oN aot ae Saat lth, ate ere etctat altel tettets)Pevict a ke seh aisle allel she’ eirsitelyolra,e 18

Second visible tergite of abdomen with quite strong apical bristles on entire margin;
apex of outer forceps of male hypopygium slightly but distinctly dilated or

SpatulateinGhext=fiune slid») mesa cee ernie ae tei hel Aedehs Chel elieelet = accepta, n. sp.
Second visible abdominal tergite without well developed bristles on hind margin
centrally; apex of outer forceps of male hypopygium acute ............ 18A
Inner, or superior, hypopygial forceps of male with a deep notch basad of middle
when seen from the side (Text-figure 12) ............-------eeees clarki, n. sp.
Inner, or superior, hypopygial forceps of male not notched when seen from the side,
straight or slightly curved ......... eee eee eee cee tt tee eee ee tees 18B

Both pairs of hypopygial forceps of male almost straight (Text-figure 14) ......
REE eT Rr aera ee eGR Wie stuntman haere ters ctoter au caleisyeNen si avieis kstatensns metallica, n. sp.
Both pairs of hypopygial forceps of male distinctly curved (Text-figure 15) ......
apne ee MeN rato ae a raretnurat Nave sb en eifcrves cilalishisinente )s\aclistioienal sietsileeltcRetfeR sts assimilis, n. sp.

Abdomen bronzy black, with dense brassy or golden dust which is distinctly
checkered; wings quite noticeably yellow at bases; prosternal hairs and
those on centre of propleura yellow; posterior sublateral bristle strong; fourth

vein distinctly curved beyond subapical angle ...........-. auriventris, nN. SD.
Abdomen metallic blue, blue-green, or bronzy, with or without whitish dusting, some-
times checkered, but never with brassy or golden dust ............-+-+-+++-- 20
Fourth vein beyond preapical angle very distinctly curved, not slightly and regularly
EROS, bb S06 6 Ho rosclolAn Blolale’oud 6 old p couse olgio dia dold/oNy Ola'o O:0'0)0.010.010-0/0.0.000'0: 0/0016 O10 21
Fourth vein beyond preapical angle straight or almost so, at most with a very
slight and regular arcuation on entire length ..............+2eeseeeeeeeee 22
Calyptrae slightly browned, centrally at least; abdomen bright metallic greenish
blue, with changeable white dusting ............-.+-+++eeeeees metallica, n. sp.
Calyptrae pure white; abdomen bronzy or greenish black, with quite dense white
dusting which is distinctly checkered ..........----++sseeeees plebeia, n. sp.

Parafacials quite densely and evenly yellowish white dusted; abdomen bluish black,
with grey dust on tergites, which is rather dense, and distinctly checkered ....
eee Reem MS PE SENDS A RORY hb S 23 (SPP Cae ets Mobo tofene A eticre koh on eulertou er eh swe slletseucltenav's minor, Nn. Sp.

Parafacials with checkered or variegated dusting, most conspicuous opposite second
antennal segment when seen from the side; abdomen bright metallic blue-green,
with rather slight whitish dusting which is most distinct laterally accepta, n. sp.

It must be noted that there are probably many species in Australia which are

not included in the above key and that the most reliable characters for the
recognition of those listed above are those of the hypopygia of the males.. The
females do not present many outstanding characters and it would be unwise to
place too much reliance upon the diagnostic characters used for this sex without
having male specimens for checking up. As already stated, the above is

308 NOTES ON AUSTRALIAN DIPTERA, Xi,

presented merely as a preliminary attempt to distinguish the native species, and
must be revised with the accession of more material, and the results checked
by rearings in the field.

CALLIPHORA OCHRACEA Schiner.

Reise Novara, Dipt., p. 307, 1868; ? Ochromyia hyalipennis Macq., Mém. Soc.
Sci. Lille, 1850 (nec 1834), p. 218; Dipt. Hxot., Suppl. 4, p. 245, Pl. xxii, fig. 10,
1850; ? Adiochosia hyalipennis Surcouf, Nouv. Arch. Mus. Hist. Nat. Paris (5),
p. 85, 1920.

This strikingly distinct species was cited as the genotype of Neocalliphora
by Brauer and von Bergenstamm. I cannot admit its claim to even subgeneric
separation from Calliphora.

I am inclined to consider this, which is the mainland form, as probably
entitled to specific distinction from the one with the dark thoracic dorsum herein
listed as a variety, but my material is so scanty that it is impossible for me to
arrive at a definite conclusion on the matter. The females before me appear to be
a little more robust, with the anterior thoracic width greater, than the Tasmanian
form.

Localities: Eungella, 45 miles west of Mackay, Queensland, 1,400-2,000 feet,
13-25 Sept., 1923 (Goldfinch); Sydney, N.S.W., 15 Sept., 1923; Dinner Creek,
Ravenshoe, N.Q. (Sherrin).

CALLIPHORA OCHRACEA, form NIGRITHORAX NOV.

This is the Tasmanian form, which differs as stated in the key and above
paragraph. Whether or not this is hyalipennis Macquart is immaterial as the
latter name is a homonym, a fact previously pointed out by Hardy.

Length, 10-11 mm.

Type, Mangalore, Tasmania, 15 March, 1914.

CALLIPHORA TIBIALIS Macquart.

Mém. Soc. Sci. Lille, 1844, p. 323 (1846); Dipt. Exot., Suppl. 1, p. 195, 1846.

Apparently one of the commonest and most widely distributed species in
eastern Australia. Readily distinguished from any other species in the genus by
the reddish tibiae, entirely pollinose abdominal dorsum, and the lack of yellow
hairs on abdomen and pleura. In the colour of the pollen on abdomen it resembles
auriventris described on a subsequent page of this paper. Quite variable in size,
running from 6 to 11 mm. in length.

Many specimens from New South Wales.

I believe that Calliphora tessellata' Macquart is the same species as tibialis.

CALLIPHORA STYGIA (Fabricius). (Text-figure 5.)
Spec. Ins., ii, p. 438, 1774.

I do not know if Patton made certain when he examined the type of this
species whether the eye facets in that specimen are enlarged or not, but accept
his assignment of specific names merely as a tentative course, definite decision
being dependent upon a careful examination of types. It appears highly probable
to me that both species may have been described, as they are widely distributed,

1Mém. Soc. Sci. Lille, 1842, p. 287 (1843); Dipt. Hxot., ii, pt. 3, p. 130, pl. xv, fig. 6,
1843.

BY J. R. MALLOCH. 309

and there are several so-called synonyms listed under stygia. In the meantime
the names accepted herein may be used.

Hardy has expressed the opinion that hilli Patton is an aberration of stygia,
and notes that “both forms are to be bred in Brisbane under conditions that point
to the progeny being from the same parent ....I have not been able to ratify
or refute this opinion ...”. In this connection it appears pertinent to indicate
that it is unwise to conclude that only one species of this genus is present in lots
reared from even a limited amount of food, several species of this and allied genera
often occurring in the same pabulum.

Dissection of the males of the two forms points to the fact that they are
distinct species. The male hypopygium of stygia is figured herein (Text-fig. 5).

Localities: Sydney, Kuringgai, Blue Mts., Kosciusko, N.S.W.; and Ruapehu,
New Zealand.

Text-figure 5. Hypopygium of Calliphora stygia. a, side view; b, upper
forceps from behind.

Text-figure 6. Hypopygium of Calliphora hilli. a, side view; b, both
forceps from behind.

CALLIPHORA HILLI Patton. (Text-figure 6.)
Phil. Jour. Sci., 1925, xxvii, p. 400.

As indicated under the preceding species this one probably has had some other
name given to it before Patton described it. The characters listed in the key
readily distinguish it from stygia as herein accepted, and the hypopygium is quite
distinct from that of its ally (Text-fig. 6). I have before me two females from
EHungella, Queensland, which have three pairs of presutural acrostichals, but they
have the narrow frons and orbits of hilli, and evidently belong here, a conclusion
strengthened by the fact that a male of hilli was taken along with them. I
dissected the male to make certain of its identity.

Localities: Sydney; Barrington Tops, Dec., 1921 (G. Goldfinch), N.S.W.;
Hungella, 45 miles west of Mackay, Queensland (Goldfinch).

The records may point to this being a more northern form but I have before
me females from New Zealand that are referable here, and Hardy records it from
Tasmania.

CALLIPHORA FUSCOFEMORATA, DN. SD.

Male.—Head fuscous, lower portion of parafacials, anterior part of cheek
beyond the raised area, lower part of face, the palpi, and antennae, brownish
yellow, third antennal segment darkened apically; occiput, raised part of cheeks,
and the orbits, grey dusted, a paler patch of dust opposite bases of antennae on
parafacial; hairs on lower half of occiput and on posterior half of cheeks yellow.

310 NOTES ON AUSTRALIAN DIPTERA, Xi,

Thorax black, with slight grey dusting, the dorsum faintly vittate, when seen from
the side the postsutural area has a narrow central dark vitta, and on each side
between the acrostichals and dorsocentrals a small dark spot at middle; many of
the pleural hairs yellow, but those on mesopleura anterior to the bristles dark.
Abdomen honey yellow, with a broad violet-blue patch on middle of basal three
tergites which is not sharply outlined laterally, the fourth tergite densely
yellowish grey dusted, with distinct checkering, the dust obscuring the ground
colour; most of the ventral hairs and bristles yellow. Coxae and femora fuscous,
the latter yellow at bases of hind pair; tibiae honey yellow, hind pair slightly
darkened at bases; tarsi black; bristles at bases of ventral surfaces of hind
femora yellow, the others black. Wings brownish hyaline. Calyptrae pale brown.
Halteres yellow.

Frons at narrowest point about as wide as anterior ocellus; parafacial about
as wide as third antennal segment; hairs on facial ridges extending to above
middle. Posterior sublateral bristle lacking. Abdomen robust. Legs normal.
Apical section of fourth vein beyond angle distinctly curved.

Length, 7 mm.

Type, Kuranda, Queensland (F. P. Dodd).

CALLIPHORA AUGUR (Fabricius).

Ent. Syst., iv, p. 321, 1775; oceaniae Rob.-Desv., Essai Myod., p. 4388, 1830;
villosa Rob.-Desv., id., p. 4387; lateralis Macq., Mém. Soc. Sci. Lille, 1842, p. 291
(1843); Dipt. Exot., ii, pt. 3, p. 184, 1843; rufiventris Macq., Mém. Soc. Sci. Lille,
1846, p. 98 (1847); Dipt. Hxot., Suppl. 2, p. 82, Pl. v, fig. 5, 1847.

This species differs from the preceding one in having the head much paler,
the lower half of occiput, entire cheeks, lower half of parafacials, and the face being
orange yellow. The pleura are quite densely yellowish dusted and appear paler
than the dorsum, the humeral angles are yellowish, the dorsum is whitish dusted
and on it there is no trace of the spot between the postsutural acrostichals and
dorsocentrals. The dorsocentral blue mark on abdomen is quite sharply limited
laterally. Coxae, femora, and tibiae orange yellow, tarsi black, ventral bristles
on hind femora of male black.

Structurally similar to the preceding species, the posterior sublateral bristle
usually lacking; male hypopygia not dissected in either species.

Length, 8-11 mm.

I have before me specimens of both sexes from Sydney, N.S.W.

Two females without more definite locality than Australia differ from those of
typical augur in having the dorsal blue mark on abdomen much broader, the
third tergite being almost entirely blue dorsally, and the fourth is entirely blue
with only slight whitish dusting which does not obscure the ground colour.
These may belong to a distinct species, but without males of each for dissection
I cannot give a definite decision either way so leave them temporarily as augur.

Patton lists a number of synonyms of augur in his recent paper and it appears
probable that further examination of the types of the species so listed may result
in the discovery that they do not really all belong to the same species. Mean-
while the only course open to me is to accept his findings. The following two
species belong to the same group as the preceding two.

CALLIPHORA MACLEAYI, Nl. SD.
Male and female.—Head coloured as in fuscofemorata, but the cheeks not so
much darkened. Thorax shining, with a violet tinge, whitish dusted, and with

BY J. R. MALLOCH. 311

mere traces of vittae on dorsum; pleura not shining and much more densely
yellowish grey dusted than dorsum, mesopleura yellow haired on lower half.
Abdomen honey yellow, broadly violet-blue on centre of dorsum of all tergites,
which colour is sharply limited laterally and widened on posterior margin of
visible tergites 1 to 3, no dense dust on fourth, the surface, like that of other
tergites, distinctly shining; most of ventral hairs and bristles yellow. Coxae,
femora, and tibiae honey yellow, tarsi black; femoral hairs and bristles yellow
basally. Wings hyaline. Calyptrae and halteres yellow.

Frons in male a little wider than in fuscofemorata, the head similar in other
respects. Thorax with three pairs of presutural acrostichals, and a distinct
posterior sublateral bristle. Otherwise similar to fuscofemorata; hypopygium not
dissected.

Length, 7-9 mm.

Type male, allotype and one male and one female paratype, Eungella, 45 miles
west of Mackay, Queensland, 1,400-2,400 feet, 13-25 Sept., 1923 (Goldfinch).

CALLIPHORA CENTRALIS, N. SD.

Male.—This species has the head coloured as augur, the cheeks being entirely
yellow, but the third antennal segment is paler than in that species. The thorax
is bluish black on dorsum, more metallic posteriorly, with whitish dusting, and
more noticeably vittate than the other species, especially anteriorly; the humeri
are yellow, and the pleura are quite densely yellow dusted. Abdomen honey
yellow, with a narrower blue-green stripe on basal three tergites, and none on
fourth, or only a trace in centre of the dark colour. In other respects similar to
macleayi; hypopygium not dissected.

Length, 9 mm.

Type, Eungella, 45 miles west of Mackay, Queensland, 1,400-2,400 feet,
13-25 Sept., 1923 (Goldfinch).

A female which I take to belong to this species has the thorax more densely
whitish dusted on dorsum, and the ground colour of the pleura largely yellow.

Locality: Townsville, Queensland (F. H. Taylor).

A second female agrees more closely with the male in colour of the thorax, but
has the fourth tergite overcast with a metallic blue-green tinge.

Locality: Upper Lansdowne, N.S.W., 25 Febr., 1921.

None of the species treated hereinafter have any yellow colour on either the
thorax or abdomen, and all of them have entirely black legs, forming a very
difficult group to distinguish specifically. Only a careful examination of specimens
and more especially an examination of the male genitalia will serve to identify
them in the adult stage but corroborative evidence of specific value may be found
either in the larval stages or in the habits of the forms involved.

CALLIPHORA SD.

Female.—I have some doubt as to the specific identity of this form. It may be
merely an aberrant form of one of the other species dealt with in this paper, but
I have no means of checking this up so leave it without a definite name meanwhile.

In colour it closely resembles clarki and accepta, with the strongest possibility
_.that it belongs to the latter.

Locality: Botany Bay, N.S.W. (H. Petersen).

312 NOTES ON AUSTRALIAN DIPTERA, Xi,

CALLIPHORA DISPAR Macquart.

Mém. Soc. Sci. Lille, 1844, p. 323 (1846); Dipt. Exot., Suppl. 1, p. 195, 1846;
C. pubescens Macq., Mém. Soc. Sci. Lille, 1850, p. 215 (1851); Dipt. Hxot., Suppl. 4,
p. 242.

I assign to this species a female specimen which is somewhat similar to the
one mentioned above, but it is fully 10 mm. in length, has the thorax more
noticeably whitish dusted and less evidently -vittate, the abdomen brassy green
instead of metallic blue, and quite conspicuously, though not densely, white
dusted and slightly checkered; the bases of wings and calyptrae yellowish.

Locality: Uralla, N.S.W.

This is the only species before me that appears to agree with Patton’s
interpretation of dispar Macquart.

CALLIPHORA APICALIS, D. Sp.

Male—Head fuscous; interfrontalia velvety black, orbits brownish grey
dusted, the dusting extending over parafacials to below middle of eye and
presenting no conspicuous spot opposite bases of antennae; lower margin of face
and the cheek at vibrissal angle rufous, raised part of cheek fuscous, with grey
dust; antennae black, apex of second and extreme base of third segment reddish;
palpi orange; hairs on centre of occiput and hind margin of cheeks golden
yellow, the others black. Thorax black, with a blue cast, dorsum slightly white
dusted and very faintly vittate. Abdomen dark metallic blue, with whitish dusting
on basal three tergites which is densest laterally and leaves a narrow dark
dorsocentral vitta, fourth tergite so densely yellow dusted that the ground colour
is hidden, and the segment is conspicuously differentiated from the others. Legs
black. Wings greyish hyaline, not yellow at bases, both basal costal scales black.
Calyptrae brown, fringes yellowish. Halteres yellow. Coverings of both spiracles
brownish yellow.

Head as in erythrocephala Meigen; narrowest part of frons about as wide as
third antennal segment; eye facets slightly enlarged above; parafacial as wide
as third antennal segment. Thorax with the bristling as in erythrocephala, the
posterior sublateral bristle strong; a fine additional notopleural bristle present in
the type. Apical bristles on all except first visible tergite long but fine, the disc
of fourth tergite with numerous long fine bristles; hypopygium not dissected, but
exposed in type so that the paired forceps are seen to be similar to those of hilii.
Legs as in erythrocephala. Apical section of fourth vein beyond angle distinctly
bent.

Length, 9-5 mm.

Type, Kosciusko, N.S.W., 7 Dec., 1922 (Goldfinch).

oa

Text-figure 7. Hypopygium of Calliphora robusta. a, side view; b, forceps
from behind.

BY J. R. MALLOCH. 313

CALLIPHORA ROBUSTA, 1. Sp. (Text-figure 7.)

Male.—The type specimen of this species is more robust than apicalis, with
a more rotund abdomen, and is not so brightly coloured, the thorax being almost
black, slightly obscured by whitish dusting, and the abdomen is greenish black on
all tergites, with slight even whitish dusting which is most conspicuous laterally.
The wings are yellowish at bases.

The area between the raised portion of cheek and the vibrissal angle in type
has numerous hairs on it while in apicalis this portion is bare. Posterior sub-
lateral bristle present but weak; only two notopleural bristles present. Hypopygium
as Text-fig. 7. In other respects similar to apicalis except that the abdominal
hairs and bristles are much shorter and weaker.

Length, 10 mm.

Type, Kosciusko, N.S.W., 7 Dec., 1922 (Goldfinch).

CALLIPHORA XANTHOCERA, n. Sp. (Text-figure 8.)

Male.—Head black, orbits white dusted, a large white dusted mark on
parafacial opposite basal half of antenna, area between vibrissal angle and raised
part of the cheek, as well as lower margin of face, rufous; basal two antennal
segments fuscous, second segment rufous at apex, third bright orange coloured,
extreme apex browned; palpi orange coloured; central part of occiput with
yellowish hairs, the other hairs on head black. Thorax as in erythrocephala, the
fine hairs black. Abdomen quite brilliant metallic blue-green, like that of a
Lucilia, with slight whitish dusting on the incurved lateral portions of the
tergites. Legs black. Wings greyish hyaline, darker at bases; both basal costal
scales fuscous. Calyptrae dark brown, the fringes pale. Halteres brownish.

' Text-figure 8. Hypopygium of Calliphora wxanthocera. a, forceps from
behind; b, forceps from the side; c, apex of penis from below; d, apex of penis
from the side.

Frons as in erythrocephala; parafacial about as wide as third antennal
segment, the latter shorter and wider than in most related species; arista haired
on little more than the basal half. All sublateral bristles of thorax long; an
additional bristle between the normal two notopleurals; scutellum with about
twelve bristles. Abdomen with the hairs on second visible tergite dense except
on middle, those on third much sparser, longer, and stronger, the apical bristles
on this tergite long and rather strong, fourth tergite with long setulose hairs
on disc which are mostly almost as long as the apical bristles; ventral hairs
longest on second and third sternites; hypopygium as Text-fig. 8. Fore tibia with
two median posterior bristles; mid tibia with from 3 to 5 anterodorsal and
posterodorsal bristles. Apical section of fourth vein distinctly curved beyond
the angle. :

314 NOTES ON AUSTRALIAN DIPTERA, Xi,

Length, 7 mm.

Type, Kosciusko, N.S.W., 5 Dec., 1921.

A female before me differs from the above type in having the third antennal
segment narrower and much more extensively blackened, the fore tibia with but
one median posterior bristle, and the fourth vein straighter on apical section
beyond the angle. It may really belong to this species as it has the additional
bristle between the two notopleurals, and otherwise closely resembles the male.

Length, 6-5 mm.

Locality: Kosciusko, N.S.W., 7 Dec., 1922 (Goldfinch).

CALLIPHORA AUSTRALICA, n. sp. (Text-figure 9.)

Male and female.—This species closely resembles the preceding one but is
larger, has the third antennal segment mostly black, lacks the additional noto-

Text-figure 9. Hypopygium of Calliphora australica. a, side view; b,
forceps from behind.

pleural bristle, and has the hypopygium as Text-fig. 9. The colour of the abdomen
is the same in both species, which distinguishes them from robusta and most
other species of the genus. Fore tibia with but one median posterior bristle.
Length, 9-11 mm.
Type and allotype,. Kosciusko, N.S.W., 7 Dec., 1922, and 14 Mar., 1920, 5,000
feet, respectively; male paratype, Blackheath, N.S.W., 24 Dec., 1921; female
paratypes, Fish River, N.S.W., 25 Mar., 1923, and Blue Mts., N.S.W., 28 March, 1923.

CALLIPHORA ERYTHROCEPHALA (Meigen).
Syst. Beschr. Hur. Zweifl. Ins., v, p. 62, 1826.

This species, like Musca domestica Linné, appears to have an almost cosmo-
politan distribution. It is readily distinguished from any other species of the
genus in Australia by its red cheeks, which are black haired except behind, and .
the dark coloured calyptrae with their white margins and fringes.

Locality: Sydney, N.S.W., 30 May, 1921. I record only the material before me.

CALLIPHORA MINOR, n. sp. (Text-figure 10.)

Male and female.—very similar to accepta in colour, but the thoracic dorsum
is more evidently vittate, and the abdomen is quite distinctly white dusted,
especially in the female where there are distinct checkerings, and each hair and
bristle is inserted in a minute black dot.

Structurally similar to accepta, but the male has the outer hypopygial forceps
as Text-fig. 10, the frons is at narrowest part about twice as wide as anterior

BY J. R. MALLOCH. . 315

ocellus, the posterior sublateral bristle is short, and the size is less.

Length, 3-5-4-5 mm.

Type, male, Townsville, N. Queensland (G. F. Hill); allotype and two female
paratypes, Darwin (G. F. Hill); one male and four female paratypes, Cairns,
N. Queensland (J. F. Illingworth). The last series in United States National
Museum.

Text-figure 10. Hypopygium of Calliphora minor, outer forceps from side.
Text-figure 11. Hypopygium of Calliphora plebeia. a, side view; b, forceps
from behind.

CALLIPHORA PLEBEIA, nh. Sp. (Text-figure 11.)

Male.—A black species, with the antennae in variable proportion reddish
yellow, sometimes entirely so; palpi rufous yellow; orbits, parafacials, and cheeks
densely yellowish grey dusted; thoracic dorsum slightly bronzy, with quite evident
whitish dust, and rather distinctly vittate; abdomen bronzy or greenish black,
with silvery white dusting, which is distinctly checkered. Wings hyaline. Legs
black. Calyptrae whitish. Halteres yellow.

The eyes are much closer together than in any other species in this segregate,
and the facets are quite noticeably enlarged on upper anterior part. Posterior
sublateral. bristle present. Bristles on apex of second abdominal segment strong
on sides only, those at apex of third stronger than those at apex of fourth, the
discal bristles on latter long, but weak and hair-like. Fore tibia with one median
posterior bristle. Hypopygium as Text-fig. 11. Apical section of fourth vein bent
beyond angle.

Length, 7-10 mm.

Type and four paratypes, Cairns, N. Queensland (J. F. Illingworth). One
labelled ‘flowers’, the others “ex corn’.

CALLIPHORA AURIVENTRIS, 0. Sp.

Female.—Very similar to tibialis Macquart. Frontal orbits and upper part of
parafacials with yellow dust; antennae and palpi testaceous yellow, third segment
of former slightly browned; cheeks fuscous on raised part, with yellowish grey
dust and black hairs. Thorax black, with whitish dusting on dorsum, and the
usual vittae almost indistinguishable; pleura coloured as dorsum, many of the
fine hairs yellow, especially on prosternum, centre of propleura, round stigmatal
bristle, on pteropleura, and hind and lower portions of sternopleura. Abdomen
black, with a bronzy cast, but the dorsum almost totally obscured by dense brassy
or golden dusting which is irregularly checkered or variegated, the hairs black.
Legs black. Wings quite conspicuously yellow at bases. Calyptrae and halteres
orange-yellow.

N

316 NOTES ON AUSTRALIAN DIPTERA, Xi,

Head normal. Thoracic dorsum with the surface hairs very fine and short;
posterior sublateral bristle quite prominent; acrostichals 2 + 3; both pairs of
intra-alar bristles strong. Legs normal. Fourth wing vein quite evenly curved
beyond angle.

Length, 11 mm. ;

Type, Fish River, N.S.W., 25 Mar., 1923.

The colour of the abdomen is very similar to that of tibialis, but the legs are
entirely black, and the pleural hairs are largely yellow, which is not the case in
that species.

CALLIPHORA CLARKI, n. sp. (Text-figure 12.)

Male.—A species with much the same habitus as some of the European forms
at present placed in Onesia. Head black, orbits white dusted, when seen from the
side the parafacial is black at base of antenna, has a large white dusted spot
above middle, and is blackish below; cheeks and occiput yellowish grey dusted;
base of third antennal ‘segment broadly rufous; palpi rufous. Thorax bluish
black, slightly white dusted, and distinctly vittate, in type with a dark line or
vitta connecting the pair in each series of presutural dorsocentrals when seen
from the side and behind. Abdomen quite conspicuously metallic green, but with
distinct white dusting which is almost evenly distributed on dorsum. Legs black.
Wings smoky hyaline, veins dark. Calyptrae brownish, margin of lower one
yellowish. Halteres brownish yellow.

Narrowest part of frons over twice as wide as anterior ocellus; eye facets
slightly enlarged on upper half anteriorly; parafacial distinctly wider than third
antennal segment; vibrissal: angle produced. All three sublateral bristles distinct.
Abdomen broadly ovate; hypopygium as Text-fig. 12. Fore tibia with two posterior
median bristles; posterior surface of fore femur longer haired than usual. Apical
section of fourth vein distinctly curved beyond angle.

Length, 7 mm. :

Type, Perth, Western Australia, 1917 (J. Clark).

Text-figure 12. Hypopygium of Calliphora clarki. a, side view; b, forceps
from behind.

CALLIPHORA ACCEPTA, nh. Sp. (Text-figure 13.)

Male and female.—Similar to clarki, differing essentially as noted in the key
to species. Hypopygium of male as in Text-fig. 13. The males have usually two
posterior bristles near middle of fore tibia, but in the females I have before me
there is but one such bristle. The abdomen is but slightly white dusted on dorsum
and the species has more the habitus of a Lucilia than a Calliphora.

Length, 6-7:°5 mm.

BY J. R. MALLOCH. 317

Type, male, allotype, one male and one female paratype, Botany Bay, N.S.W.
(H. Petersen); one male paratype, Como, N.S.W., December, 1923, swept from
flowers (H. Petersen); two male and one female paratypes, Blue Mts., N.S.W.;
one male paratype, Mooroopna, Victoria, 31 Dec., 1923 (F. E. Wilson); one male
paratype, Sydney, N.S.W., 16 Oct., 1921.

Text-figure 13. Hypopygium of Calliphora accepta. a, side view; b,
forceps from behind.

CALLIPHORA METALLICA, n. sp. (Text-figure 14.)

Male.—A larger and more robust species than the preceding one, with the
abdomen more noticeably white dusted, and the second abdominal tergite without
the well developed central bristles on hind margin. The hypopygium is as shown
in Text-fig. 14, but in other respects the two species are very similar.

Length, 9-10 mm.

Type, Woolgoolga, N.S.W., 27 Jan., 1923; paratype, Orange, N.S.W., 21 Apr.,
1923.

CALLIPHORA ASSIMILIS, n. sp. (Text-figure 15.) _
Male and female.—More like plebeia in colour and habitus than like the
preceding species, but the eyes have the facets but little enlarged on upper half,
the frons is wider in male, and the parafacials.are not so evenly and densely
dusted. The hypopygium of male is as shown in Text-fig. 15.
Length, 8-9 mm.

Text-figure 14. Hypopygium of Calliphora metallica, side view.
Text-figure 15. Hypopygium of Calliphora assimilis, side view.

Type, allotype, and one male paratype, Marwood, near Mackay, Queensland,
Jan., 1924 (W. C. Harvey).

It must be remembered that in my opinion there are more species belonging
to this last section than are included in the above key and descriptions. Colour

318 NOTES ON AUSTRALIAN DIPTERA, Xi,

descriptions are almost valueless here and only very careful comparative work
and hypopygial examinations will disclose just how many there really are in
Australia, facts which are impossible of my ascertaining from the small amount
of -material now in my hands.

Text-figure 16. Head of Pollenia hirticeps, side view.
Text-figure 17. Apex of wing of Pollenia hirticeps.
Text-figure 18. Hypopygium of Pollenia hirticeps, side view.

Genus PoLLENIA Robineau-Desvoidy (1830).

This genus is represented in Australia and New Zealand by species which
differ from the genotype, rudis Fabricius, in having the extreme basal portion of
the stem vein on under side without evident hairs apically. This segregate if
entitled to generic rank will bear the name Sepimentum Hutton, but I consider it
as at best entitled to subgeneric rank.

The head (Text-fig. 16) is quite different from that of typical species of
Calliphora and the lower calypter is bare. The supraspiracular ridge and centre
of propleura are bare, characters which distinguish the Australian species from
those of Lucilia. In the Australian species of Pollenia the prosternum is also bare.
The venation of the apex of wing is quite different from that of Anthracomyia
(Text-fig. 17). There is but one Australian species before me.

POLLENIA (SEPIMENTUM) HIRTICEPS, n. sp. (Text-figures 16, 17 and 18.)

Male and female.—Head black, grey dusted on orbits, raised part of cheeks,
and upper parafacials, the anterior part of cheek and lower part of parafacial
reddish, and not grey dusted; antennae reddish testaceous, third segment more
or less browned apically; arista black; palpi coloured as antennae; hairs on
frons, parafacials, and cheeks, black, some of those on lower occiput and a few
on hind part of cheek yellow. Thorax black, with a bluish cast, the dorsum thinly
whitish dusted and almost entirely without vittae; dorsal hairs all-black, the
curled or crinkly yellow hairs confined to parts of pleura. Abdomen variable in
colour, black with cupreous or blue lustre, or metallic bluish black, with thin
regular whitish dusting on dorsum and no markings, the hairs all black. Legs
black. Wings slightly smoky, darkest at bases. Calyptrae and halteres brown.

Male.—Head in profile as in Text-fig. 16; frons at narrowest point not wider
than third antennal segment, the anterior half forming a large triangle, orbital
hairs long and strong, in several series, those on the parafacials also long and
curved, the entire vibrissal area sparsely short haired; antennae distinctly
separated at base, but no carina on face; longest hairs on arista not longer than
width of third antennal segment, the latter rather narrow. Thorax with 2 + 3

BY J. R.. MALLOCH. : 319

dorsocentrals, 2 + 2 acrostiehals, two pairs of intra-alars, the prealar long, two
bristles on the presutural area, and two sternopleurals. Abdomen narrowly ovate;
hypopygium as in Text-fig. 18. Legs as in Calliphora erythrocephala Meigen.

Female.—Frons at vertex one-third of the head width, much widened to
anterior margin, orbits with a series of quite long, strong, incurved bristles along
the inner margin, several series of shorter forwardly directed setulae laterad of
these, a quite prominent forwardly and outwardly directed bristle just in front of
level of anterior ocellus, and an outwardly directed one between it and the
vertical bristles. Abdominal tergal bristles weak.

Length, 6-8 mm.

Type, male, allotype, and one female paratype, Blue Mts., N.S.W.; paratype
males, Blackheath, N.S.W., and Mt. Lofty Ranges, S. Aust.

Genus ANTHRACOMYIA, novum.

This genus is rather similar to Pollenia, but there are no soft curled or
crinkly hairs on thorax, and the venation of the wing is quite. different apically
(Text-fig. 19). Genotype, A. atratula, n. sp.

ANTHRACOMYIA ATRATULA, 0. Sp. (Text-figure 19.)

Male.—Black, shining. Face, parafacials, and raised part of cheeks, whitish
dusted; antennae and palpi black; cephalic hairs all black. Thorax slightly
whitish dusted on dorsum, without evident vittae. Abdomen glossy black, when
seen from behind with slight, even, whitish dusting and a trace of a dark central
line basally. Legs black. Calyptrae fuscous. Halteres brownish yellow.

Frons linear above, forming a triangle on anterior half, orbits on widened part
of frons gradually widening anteriorly, with a series of long setulose hairs
along inner margins and a few fine short hairs laterad of these; parafacial about
as wide as third antennal segment, with rather long hairs almost to lower margin
of eye; the cheek adjacent to vibrissal angle almost bare; one or two hairs above
vibrissae; arista haired almost to apex, the longest hairs about as long as width of
third antennal segment; palpi slender. Thorax with 2 + 3 dorsocentrals and
apparently the same number of acrostichals, two pairs of long intra-alars, a long
prealar, two bristles on presutural area, no upper anterior bristle on mesopleura,
and, like the preceding genus, no hairs on the lower anterior half of pteropleura.
Abdomen narrowly ovate, the dorsal hairs rather sparse, long and setulose. Legs
as in last species, the ventral median bristle present on mid tibia; fore legs in
type missing. Wing venation as in Text-fig. 19.

Length, 4 mm.

Type, Allowrie, Killara, N.S.W., 30 Jan., 1921 (Waterhouse).

Text-figure 19. Wing of Anthracomyia atratula.
Text-figure 20. Hypopygium of Chrysomyia megacephala a, side
view; b, penis from above.

320 NOTES ON AUSTRALIAN DIPTERA, Xi,

This small species is represented by a male in damaged condition, but it is
so clearly distinct from any other species known to me that I have decided to
describe it. The character of hairs on the lower anterior half of pteropleura is
quite an important one in this group, and may be used to distinguish doubtful
Specimens of small species of Calliphora in which the hairs on the upper surface
of lower calypter are not readily distinguishable from this and the preceding genus,
the hairs being always present on lower anterior half of pteropleura in Calliphora.

Genus Luciz1A Robineau-Desvoidy (1830).

This genus is of considerable economic importance in Australia because of
the habits of some of the species which feed in the larval stages in the matted
wool of sheep, and penetrate the flesh. There are known to me four well
distinguished segregates of the genus, which I accept as subgenera. Three of
these I have seen from Australia, and below I append a key for their recognition,
the other occurs only in the Oriental region so far as I am at present aware. I
have published elsewhere a key to all four subgenera, but this is not readily
available to Australian students. It is worth mentioning that in many species
of the genus there are sparse microscopic hairs on the face, a character found
also in Phumosia Robineau-Desvoidy, but in one or two species of the latter the
hairs are quite prominent.

Key to Australian Subgenera.
1. The convexity immediately above metathoracic spiracle with conspicuous fine erect

LOE D ESS 9S con ROME ET BCOLE Gt CES AEA RIC ERPS Oy DRT Et BER ARB eH arty Hemipyrelia Townsend

The metathoracic convexity without erect fine hairs ................0-ceeeeees 2

2. Sclerite at extreme base of stem vein on under side of wing with short setulose hairs
DICE iy seat cce fase tec seis, aries samen sean lia salen Wear atictse la seits apes tee eee Lucilia Robineau-Desvoidy

Sclerite at extreme base of stem vein on under side of wing without evident hairs
ShanvdooadcDOOODOUAUOO eA Od HOMO oO MOO.o auo abo. Phenicia Robineau-Desvoidy

Subgenus HEMIPYRELLIA Townsend (1918).
In addition to having erect fine hairs on the metathoracic convexity this
subgenus has setulose hairs on the basal sclerite of stem vein below.
I have seen five species of the subgenus, two of which occur in Australia.

LuciLiA (HEMIPYRELLIA) FERGUSONI Patton.
Phil. Jour. Sci., 1925, xxvii, p. 403.

This is the only species of the subgenus known to me in which there is any
yellow colour on the abdomen, and in which the legs are not entirely black. It
bears a striking superficial resemblance to Chrysomyia incisuralis Macquart, but
is a true Hemipyrellia. Patton’s description is reliable as to colour, but the
essential characters for the recognition of its subgeneric status are not mentioned
by him. The setulose hairs on basal sclerite of first vein below are yellow, which
is unique, as is also the yellow colour of the head, pleura, and most of the abdomen.

Localities: Sydney, N.S.W., April, October, and November; Eungella, 45 miles
west of Mackay, Queensland, 1,400-2,000 feet, 13-25 Sept., 1923 (Goldfinch).

LuciniA (HEMIPYRELLIA) CYANEOMARGINATA Macquart.
Mém. Soc. Sci. Lille, 1850, p. 221, (1851); Dipt. Hxot., Suppl. 4, p. 248.
I have identified as this species one which occurs commonly in the Philippines,
Straits Settlements, and Federated Malay States. I have now before me a male and
female from Queensland.

BY J. R. MALLOCH. 321

It should be noted that Patton records under Lucilia nosocomiorum Doleschall
a number of synonyms including cyaneomarginata Bigot. Bigot described a
cyaneocincta, but I can find no cyaneomarginata Bigot. I have accepted noso-
comiorum as the name for a larger species which is apparently the one identified
as such by Patton, but it belongs to the subgenus ZLucilia.

In cyaneomarginata the frons of the male is distinctly wider than third
antennal segment, the thorax has two pairs of postsutural acrostichals, and the
abdomen has conspicuous dark blue apices to the tergites. Length, 7-8-5 mm.

Localities: Cairns, N. Queensland, carrion (J. F. Illingworth).

Subgenus PHENIcIA Robineau-Desvoidy (1830).

The members of this subgenus known to me have the eyes of the males more
widely separated than is the rule in the other two subgenera occurring in
Australia, and the basal sclerite of stem vein is bare.

There are two species known to me, both of which I have seen from Australia.

LUCILIA (PHENICIA) SERICATA Meigen.
Syst. Beschr. Hur. Zweifl. Ins., v, p. 53, 1826.

This bright metallic blue-green species has the ventral surface of abdomen in
male only moderately setulose, and in both sexes the dorsum of abdomen is
without conspicuous white dusting.

I have before me many examples from Australia, some labelled, ‘““Bred from
Wool.”

LUCILIA (PHENICIA) CUPRINA Wiedemann.

Aussereurop. Zweifl. Ins., ii, p. 654, 1830; pallifrons Bigot, Ann. Soc. Ent.
France, 1877, p. 257.

A coppery or bronzy coloured species with very noticeable whitish dusting on
thorax and abdomen. The ventral surface of abdomen in the male is furnished with
quite dense long conspicuous bristles.

I identify as this species a female from Kuranda, Queensland (F. P. Dodd).

Subgenus Lucint1a Robineau-Desvoidy.

This subgenus is represented in the material before me by two rather aberrant
species. The genotype, caesar (Linné) has quite numerous fine hairs on central
part of propleura, while the other two have no such hairs or there are but one
or two such on anterior margin. The face in some of the species is usually bare,
while in others it is invariably furnished with short sparse hairs in middle.

I present below a key for the identification of the two species I have seen
from Australia and include caesar, which I have seen no Australian specimens of
so far.

Key to Species.
1. Face invariably with some microscopic black hairs:centrally from lower to upper

extremity ; abdomen metallic blue, with dark apices to tergites; lower calypter dark
brown; centre of propleura bare or with one or two weak hairs on anterior

TUVALA EE aecist en Sicel ole yAlly, ee vias lege bode voy aces s Sled ate pevies arabe ler el otonat otis nosocomiorum Doleschall
Face bare or with some inconspicuous microscopic hairs near lower margin; abdomen
metallic blue-green or blue, without dark apices to tergites ................ 2

2. Antennae black or fuscous, third segment reddish at base; frons in female about as
wide as length of antennae; centre of propleura with quite evident pale hairs;
GCANVDUGA Sy DUT Sw WILE) > ap saisa¥ dicotgo seuspehey ey gust cues ieliotebe. at's Geol Wy ad eg onGlee aya sonst > wtaesi caesar (Linné)

Antennae orange yellow; frons in female much narrower than length of antennae;
centre of propleura bare, at most with a fine hair or two on anterior margin;
lowermcalyprenmbrownishsyellowseee ais ose ieee ee ceieens flavicornis, n. sp.

322 NOTES ON AUSTRALIAN DIPTERA, Xi,

Lucinia (LUCILIA) CAESAR Linné.
Fauna Suec., p. 451, 1761.

A brilliant metallic blue-green species which frequently has a golden tinge on
abdomen and thorax. There are no traces of dark apices to tergites in either sex.

The species has been recorded from a wide range of localities, but as a matter
of fact it is not so commonly found as the records would indicate. I have seen no
Australian specimens in either the material sent to me or in the United States
National Museum, where there is a good representation of the family from
Australia. The characters included in the above key ought to serve to separate it
from other Australian species if it does occur.

Lucit1a (Lucia) NosocomiorumM Doleschall (1857).
Nat. Tijdschr. Ned-Ind., 1857, xiv, p. 413.

Very similar to caesar, but more constantly metallic blue in colour, with dark
apices to the abdominal tergites, and usually larger in size and more robust,
averaging over 8 mm. in length.

Localities: Kuranda and Townsville, Queensland.

I can detect no outstanding differences in the structure of the Australian
specimens and some from Sumatra and the Philippine Islands. The male hypopygia
agree, and though the anterodorsal surface of the hind tibia in the Australian
males has a regular series of short setulae and there is at least one outstanding
bristle in the series on those from the Orient I consider it highly probable that
metilia Walker, which is the name given to the latter, is the same as nosocomiorum,
the latter name having priority.

Lucit1A (LUCILIA) FLAVICORNIS, N. Sp.

Female.—Frons black, slightly reddish in front, upper orbits and ocellar region
slightly shining and faintly bluish; occiput black, grey dusted, cheeks and face
orange coloured, white dusted, cheeks slightly darkened posteriorly, face darkened
above; antennae entirely orange-yellow, basal two segments slightly darker; palpi
orange. Thorax metallic violet-blue, dorsum rather evenly white dusted, the
dusting most noticeable when seen from behind, and without outstanding vittae.
Abdomen concolorous with thorax, but the dorsum on disc green, white dusted on
incuryed lateral portions of tergites. Legs dark brown or fuscous, femora with
bluish tinge, trochanters and tibiae brownish yellow. Wings greyish hyaline.
Lower calypter brownish yellow. MHalteres yellow.

Frons at vertex hardly over one-fifth of the head width, and narrower than
length of antennae; face bare. Thorax as in caesar, but the propleura is bare on
centre. In other respects as caesar.

Length, 9 mm.

Type, Cairns, N. Queensland, from dung (A. P. Dodd).

I can find no description that appears to fit the above species and though I
do not ordinarily undertake to describe species from single females in revising a
group it devolves upon one to designate all material by names which will serve
to distinguish them until final adjustments have been made.

Genus PARATRICYCLEA Villeneuve (1913).
This genus is represented in the material before me by one species which
belongs to the subgenus Caiusa Surcouf. It is possible that it has been described
by one or more of the older authors, but it is absolutely impossible to make certain

Co

BY J. R. MALLOCH. 32

of most of their species, and even after a number of attempts have been made by
recent workers to elucidate their types we are still very much in the dark as to
their exact identity, and as to even their generic positions. I consider it much
better to describe any species aS new which does not agree absolutely with the
description of any of those species and do not consider it unscientific to do so
even in cases where there is an apparent agreement if the specimens are from
different localities within the same faunal region. Only a careful examination of
types by some competent specialist will definitely settle specific identities, and
prevent the erroneous recording of species from localities in which they do
not occur.

Surcouf has recently described the chaetotaxy of Phumosia analis Macquart so
thoroughly that it is clear the species is not the same as the one described herein.

PARATRICYCLEA (CATUSA) SURCOUFI Bezzi.
Bull. Ent. Res., 1927, xvii, p. 246.

Male and female.—Shining fulvous testaceous; upper occiput, raised part of
cheeks, and frontal orbits grey or fuscous, upper part of frons in female fuscous.
Abdomen broadly blackish blue at apex. Legs fulvous yellow. Wings yellowish
hyaline. Calyptrae and halteres fulvous.

Frons in male reduced to a line, the facets much enlarged on upper half of
eye and rather sharply separated from those of lower half; frons in female a
little less than one-third of the head width, anterior supraorbital strong, inter-
frontalia bare; face normally with some microscopic hairs in centre. Thorax with
3 + 4 dorsocentral bristles, the anterior postsutural pairs short, 1 + 2 pairs of short,
widely separated, presutural, and one pair of prescutellar acrostichals; sides of
scutellum haired, the hairs on ventral surface yellow; pleural hairs yellow, those
on upper half of mesopleura black; sternopleurals 1 + 1. Superior hypopygial
forceps of male hollowed out behind, slightly curled, and tapered to apices, the
inferior as long as superior pair, stouter, and obtuse at apices. In other respects
similar to indica Surcouf, the hind tibia with two posterodorsal bristles; no hairs
at junction of humeral cross vein and subcosta, but one or two below at junction
of first and second veins.

Length, 7-8 mm.

Localities: Male, Melville Island, N.T. (G. F. Hill); Darwin, N.T. (G. F. Hill) ;
females, Stapleton, N.T. (G. F. Hill), and Marwood, near Mackay, Queensland
(W. C. Harvey); original locality, Queensland (Bezzi).

In the Oriental species before me most, or all, of the hairs are black or
fuscous, and the male has the superior hypopygial forceps more slender, not
curled, and shorter. Macquart’s species analis was described from New Holland

but I have seen no species which could be considered as possibly this from
Australia.

Genus HurpHumosiA Malloch (1926).

This genus, like the preceding one, belongs to the arbitrary group ‘Muscidae
Testacea”, and the genotype, papua Guerin, was included by Surcouf in the genus
Phumosia Robineau-Desvoidy. I recently erected the genus Euphumosia for its
reception.

In addition to the characters which may be culled from the key presented
on a preceding page of this paper it may be mentioned that the dorsocentrals are
2+ 3, acrostichals 2 + 2, intra-alars 2 pairs, and sternopleurals 1 + 1; in the male

gf

gtfn

324 NOTES ON AUSTRALIAN DIPTERA, Xi,

the frons is reduced to a mere line, the facets of the eyes are much enlarged on
upper half, and there is a strong submedian ventral bristle on mid tibia in both
sexes.

EUPHUMOSIA PAPUA (Guerin-Méneville).

Voy. Coquille, Zool., 1830, Insects, Pl. xxi, fig. 3; eristaloides Walker, Proc.
Linn. Soc. Lond., 1858, iii, p. 106; calliphoroides Walker, Proc. Linn. Soc. Lond.,
1861, v, p. 245; papouana Bigot, Bull. Soc. Zool. France, 1887, p. 610; variegata
Bigot, Bull. Soc. Zool. France, 1887, p. 610.

A testaceous yellow species, the dorsum of thorax more fulvous and with
three broad chocolate coloured vittae which are connected by a similarly wide
transverse band at their hind extremities some distance from hind margin of
mesonotum. Abdomen coloured as thorax, hind margins of all tergites deep black,
the basal one least broadly so, anterior margins of tergites 3 and 4 narrowly black,
the transverse pale fascia on each of these tergites densely whitish yellow dusted.
Legs yellow, tarsi darker. Wings browned, especially along veins.

Length, 11-12 mm.

Locality: Melville Is. (W. D. Dodd).

This conspicuously marked species has been described several times, the names
known to me including eristaloides Walker, variegata Bigot, and papouana Bigot,
while Johnston and Hardy recognize in it calliphoroides Walker.

The species does not appear to be a common one, and nothing is known of
its immature stages. There is one female in the United States National Museum.

DEXOPOLLENIA CHRYSOTHRIX Bezzi.
Bull. Ent. Res., 1927, xvii, p. 231.

This species has recently been described from N.S. Wales, but is unknown
to me.

Subfamily Chrysomyiinae.

The Australian members of this subfamily are distinguished from other sub-
families by the presence of hairs, or setulae, on the posterior upper side of the
stem vein of wing, and the haired upper surface of lower calypter. Only the
Rhiniinae have the setulae on the stem vein, and these have no hairs on the lower
calypter. In addition to these characters there are only two sternopleural bristles
present (1 + 1), the stigmatal bristle is usually present and distinct, and in many
species there are well developed hairs on the small knob-like protuberance below
and in front of the wing base.

Patton has recently dealt with the species of Microcalliphora, which he did
not separate from Chrysomyia, and he redescribed varipes Macquart and described
two new species in the same paper. He was evidently unaware of the fact that
the frons in the males of this genus is almost as wide as in the females, and his
annulipes was described, not from a female as he states, but from a male, and
undoubtedly that of varipes as it agrees in all particulars with that species. His
fulvipes had previously been described by Aldrich as flavifrons. There are thus
but two known species of the genus.

I attempt a provisional review of the species of the genera known to me
herein but the paucity of material available does not lend itself to a comprehensive
treatment of the group. It is highly probable that there are some more Australian
species yet to be discovered. There are some species of the genus which cause

BY J. R. MALLOCH. 325

myiasis in man as well as in other mammals and which ought to be guarded
against.
Below I present a synopsis of the three genera in my material now.

Key to Genera.
1. The small knob-like protuberance below and in front of wing base without any

GUUesWeoGhhoy¥= INNIS) Golo Uodonoo obo D obo conloao cotton oo olor Hucompsomyia, novum.
The small knob-like protuberance below and in front of wing base with numerous
QUESTA SIAM AITS Beier tals renee tate cos cNsrro asic; Soa ce) ale elresenel cep wuaiter onoitile (eles e eee eneheeleetatera 2

2. Eyes of male narrowly separated above, the frons almost linear ..................
BORG ESO IETS CeOSENO BCL CLD SO CLCROICRE RATER EN CRC ESOL Oa GEnTy MRE AGE CRC Reet St Chrysomyia Robineau-Desvoidy

Eyes of male widely separated above, the frons at least half as wide as either eye
and but little narrower than that of female ........ Microcalliphora Townsend

Genus EHucoMpsomy!aA, novum.

Generic characters.—Similar to the two other genera included above, the
centre of propleura, prosternum, and entire pteropleura, haired; basal sclerite
of stem vein of wing on under side haired, but, like in the American genera of the
subfamily, the knob-like protuberance below and in front of wing base is bare.
The American genera, however, have the anterior half of the pteropleura bare,
and the lower calypter in these is haired at base only, not on the entire upper
surface as in the Australian genera.

Genotype, Hucompsomyia semimetallica, n. sp.

There are two species of this genus known to me which may be distinguished
as below.

Key to Species.

Eyes of male very narrowly separated above, almost touching at middle of frons;
abdomen honey yellow, second tergite centrally, third on most of disc, and all of
fourth, metallic blue, with whitish dusting .................... semimetallica, n. sp.

Eyes of male quite widely separated above, width of frons at narrowest part about one-
sixth that of head; abdomen honey yellow, with the following bluish black markings:
First visible tergite with a narrow apical fascia, second with a moderately broad
central mark which extends along hind margin and narrows laterally into a slender
fascia, third with a still broader mark of similar nature, and fourth with a narrow
IN'COMPLELEMCEMbEBAlMEVATEA, Gye ale vatctenct cues cosucl cleus euete eu ete ual [nleselaeie lontialienencuenete latifrons, nN. sp.

EUCOMPSOMYIA SEMIMETALLICA, N. Sp.

Male.—Head fulvous yellow, hairs on cheeks, parafacials ,and lower half of
occiput, yellow. Thorax fulvous, dorsum almost entirely violet-blue, pleura
darkened on lower posterior part, metanotum blackish blue. Abdomen honey
yellow, dorsum largely violet-blue, fourth tergite more greenish, and with whitish
dusting. Legs fulvous, hind tibiae slightly browned. Wings yellowish hyaline.
Calyptrae and halteres fulvous.

Frons linear above; eye facets a little enlarged on upper half; cheek over
one-third as high as eye; palpi normal. Thorax with 2 + 4 dorsocentrals, 2 bristles
on the presutural area, one behind the humerus and the other transversely placed
from its hind margin; only one pair of prescutellar acrostichals present; sterno-
pleurals 1 + 1; anterior intra-alar short. Abdomen short ovate. ‘Tibial bristles
short, similar to those of Chrysomyia species. Apical cross vein much curved.

Length, 7 mm.

Type, Kuranda, Queensland (F. P. Dodd).

326 NOTES ON AUSTRALIAN DIPTERA, Xi,

EUCOMPSOMYIA LATIFRONS, N. SD.

Male.—Similar to the above species but the third antennal segment largely
brown, facial ridges with numerous black hairs above vibrissae which extend to
above middle; pteropleura, sternopleura, and hypopleura bluish black, with grey
dusting; legs yellow, with the apices of tarsi darkened; otherwise as stated in
the key.

The facets are but little enlarged on upper half of eye, the parafacials are
bare centrally. Thorax as in the above species, but the anterior intra-alar is absent
in type. In other respects as semimetallica except as stated in key.

Length, 7-5 mm.

Type, Loowanna, HE. Dorrigo, N.S.W., 31 Jan., 1923.

This species presents almost the same divergence from the genotype as does
Microcalliphora from Chrysomyia, but I retain them both here because of lack of
a knowledge of the characters of both sexes, and paucity of material.

Genus MICROCALLIPHORA Townsend (1916).

As stated on a previous page in this paper there are but two known species
of this genus. I have examined both of these in the United States National
Museum and give below a synopsis for their recognition.

The small size of the species and the widely separated eyes of the males
ought to readily distinguish the insects even in the field. Intensive collecting may
result in the discovery of yet unknown species of the genus. The female of
flavifrons has not been described.

Key to Species.
1, Femora and tibiae entirely yellow; frons, in male at least, blackened only on the
upper third, the anterior third bright yellow; fore femur in male with no

CONSPICUOUS Whites hairsae erase eet eee ee nee flavifrons Aldrich
Femora largely black, tibiae partly so; frons in both sexes fuscous to anterior margin;
fore femur in male with dense erect white hairs above ...... varipes Macquart

MICROCALLIPHORA FLAVIFRONS Aldrich.

Proc. U.S. Nat. Mus., 1925, lxvi, art. 18, p. 20; fulvipes, Patton, Phil. Jour. Sci.,
1925, xxvii, p. 410.

Known only from Gordonvale and Cairns, Queensland, the former the type
locality of flavifrons, the latter the type locality of fulvipes, both of the types
collected by Dr. J. F. Illingworth and sent by him to the describers.

MIcROCALLIPHORA VARIPES (Macquart).

Lucilia varipes Macq., Mém. Soc. Sci. Lille, 1850, p. 222, (1851); Dipt. Haxot.,
Suppl. 4, p. 249, Pl. xxiii, fig. 4; annulipes Patton, Phil. Jour. Sci., 1925, xxvii,
p. 410.

I have already indicated that annulipes Patton appears to me to be this
species. My specimens are from Sydney, N.S.W.; it occurs also in Queensland.

Genus CHRYSOMYIA Robineau-Desvoidy (1830).

It must be noted that the original spelling of this genus name is Chrysomya,
and that the emendation given above, which has been generally accepted, causes a
conflict with Chrysomyia Macquart, a genus of Stratiomyiidae, which was erected
at a later date, and which is generally given the name Microchrysa Loew.

BY J. R. MALLOCH. 327.

This is the most generally distributed genus of the family in the Old World,
and the one which is of most economic importance. As is usual in the case of
species that enter into economic relations with man and domestic animals there
are doubts as to the exact distribution and names of several of the species, one
of them being recorded under several names and from Africa, Asia, and Australia.
It is very desirable that accurate data be obtained to determine if it is one species
or several that occur over this territory, but I am not in possession of material
in sufficient quantities to do this work at this time. Under the species in the
text I present some points which appear to call for investigation. The following
key is given to facilitate identification of the four species which I recognize or
have reason to accept as occurring in Australia.

Key to Species.

1. Abdomen honey-yellow, with a narrow black hind marginal band on each of the
basal three visible tergites, and a broad metallic blue suffusion on dise of same
tergites; legs honey-yellow, tarsi black; hairs on the knob-like process below
base of wing golden yellow; calyptrae yellowish white; frons of female at vertex
NOtOvierrone-litthwor che! head swidth scsi. aioe cis shee ole incisuralis Macquart

Abdomen entirely metallic blue or green, with dark apices to tergites; legs black;
frons of female over one-fifth of the head width at vertex ................. 2
2. Calyptrae white or yellowish; coverings of both prothoracic and metathoracic spiracles
white; hairs on Knob-like process below wing base pale .... albiceps Wiedemann
Calyptrae fuscous or dark brown, white on outer or connecting angles; coverings
of both pairs of thoracic spiracles and hairs on knob-like process below base of
Tb? LNA. Ore: TEDISOLLISMGidiorarc| 6 cia etorotais ohoicord ol olarc ea aLO enorclon G ciated io casLomnd Ona moroIectc 3

3. Facets of upper half of eyes in male exceptionally enlarged and sharply differentiated
from those of lower half; a number of black setulae on vibrissal angle; hairs on
fourth tergite practically all black on dorsum, the pale hairs on _ sides
inconspicuous; base of scutellum darker than apex ...... megacephala Fabricius

Facets of upper half of eyes in male only moderately enlarged; no black setulae
on vibrissal angle besides the single bristle; hairs on fourth tergite practically
all white in female, in male they are mixed black and pale on disc, yellow and
CONSHICUOUSMOIM SUG SSH 5) c2rayis 15) oi estanley evel cyeusucleu syed Sheharisyek shalfeus\ leveiiepel ens teaors micropogon Bigot

CHRYSOMYIA INCISURALIS (Macquart).

Mém. Soc. Sci. Lille, 1850, p. 219 (1851); Dipt. Hxot., Suppl. 4, p. 246, 1851;
Psilostoma incisuralis Surcouf, Nowv. Arch. Mus. Nat. Paris, 1920, (5), p. 58.

Macquart originally included this species in his genus Ochromyia because of
its yellow colour, and Surcouf erected the genus Psilostoma for its reception, but
it differs but little in structure from the other species of the genus and is a
member of Chrysomyia without any doubt.

The species does not occur outside of Australia so far as I know at present.
My specimens are from Cairns and Eungella, Queensland, and Sydney, N.S.W.

CHRYSOMYIA ALBICEPS (Wiedemann).

albiceps Wiedemann, Aussereurop. Zweifl. Ins., ii, p. 404, 1830; ? rufifacies
Macq. Mém. Soc. Sci. Lille, 1850, p. 216 (1851); Dipt. Exot., Suppl. 4, p. 243.

If this identification is correct this is the most widely distributed species of
the genus, being originally described from Africa, and recorded from southern
Asia, most of the islands in the Indian and South Pacific Oceans, and Australia.
There is some question as to whether putoria Wiedemann is distinct from
albiceps, but lack of material prevents me from arriving at a definite decision on
the point. . It appears worth mentioning that there is considerable variation in
the colour of the hairs on the upper side of the lower calypter in my specimens.

328 NOTES ON AUSTRALIAN DIPTERA, Xi,

In some these are all black, in others the black hairs are confined to the central .
part of disc, while in others they are all white.

I hope to be able later to decide the matter of specific relations here if I
become possessed of sufficient material. j

I have specimens from Queensland and New South Wales.

CHRYSOMYIA MEGACEPHALA (Fabricius). (Text-figure 20.)
Ent- Syst., iv, p. 317, 1794.

This species has been recorded as occurring in Australia but so far as my
material is concerned I have not seen any specimen which agrees absolutely with:
those I have from the Orient. I have used in the foregoing key the characters
which appear to me to be of value in distinguishing it from micropogon, but I am
not at all satisfied that they will suffice for that purpose in all cases. I made a
dissection of the hypopygia of the males of two specimens which I believed were
distinct but found there were no outstanding characters in these organs by means
of which the species might be distinguished. I figure the hypopygium of the true
megacephala from the Philippines (Text-fig. 20). A comparison of this figure
with any of those of the other genera figured herein will at once show
characteristic differences.

CHRYSOMYIA MICROPOGON Bigot.
Bull. Soc. Zoo. France, 1887, xii, p. 601.

I have but one pair of this species before me, which form the basis for the
distinguishing characters given in the key and the notes in above paragraph. The
hypopygium of the male specimen was badly eaten by some pest so that only the
major features were distinguishable but they are identical with those of
megacephala.

It is not impossible that the variation in size of eye facets in this species
from that of typical megacephala is not an indication of specific distinction, but
this character is quite dependable in related groups so that I do not care to suggest
that it is not here. I have already indicated that the hypopygia of the males of
closely related species in the genus Musca are so similar that they cannot be
depended upon to identify readily the species involved, and it is quite within the
bounds of possibility that the same applies here.

Localities: Marwood, near Mackay, Queensland, Jan., 1924 (W. C. Harvey),
and Glenreagh, N.S.W., 2 Febr., 1923.

Careful field work may develop whether or not there are two species in this
complex, but laboratory findings are not satisfactory in such cases.

. Subfamily Rhiniinae.

The members of this subfamily have setulae or hairs on the basal section of
stem vein on its upper side, and the sternopleurals invariably two in number
(1 + 1). The lower calypter is usually much narrower than in Chrysomyiinae,
rounded at apex, and without hairs on its upper surface, characters which
distinguish the species from those of Chrysomyiinae.

The group is, if we except Pollenia and its nearest allies from it, exclusively
Old World in its distribution, and its centre of dispersal appears to be Africa
and the Orient, a few species overlapping into Australia and Europe. I have
recently published a key for the recognition of the genera known to me, and
include below one which includes those genera I have seen from Australia.

BY J. R. MALLOCH. 329

In this subfamily, as in the others, there are several species described by
Macquart which are difficult to identify with certainty. I have attempted to make
out these species in my material, but with what success remains to be determined
by an examination of the types, if such are in existence.

Very little is known of the immature stages of the members of the group.

Key to Australian Genera.

1. No outstanding stigmatal bristle present, the stigmatal hairs white or yellow; arista
with rather long hairs on upper side, bare below ...............-.......-- 2

One or two outstanding black stigmatal bristles present which are readily distinguished
from the pale st?gmatal hairs; arista with very short hairs both above and below

so ooooesoueboo oO DUBBO OUBDDe Ooo boo do ocladoddoaoobID Gd oIC Metallea van der Wulp

2. First posterior cell of wing closed at apex, the apical fused part of veins 3 and 4 quite
COAOMCUOWS cooddedcononooun nob od oon one ooo o oOo CmOUOnOD Chlororhinia Townsend
inst posterior cell open, or closed only in margin of wing’ ..:.......7....-5.-.. 3
Hind margin of mesopleura with but two bristles, at upper corner .................
5000000 DUO EEOC EDU UOMADE DO Ay CoG coDCODUOOdU oboe dae Rhinia Robineau-Desvoidy
Hind margin of mesopleura with a complete series of bristles ..................
PI er had cieiisis la foi'scbhs ous /a:s|lelguislia raya vamepia ta caer sbedepeicueladarsvenencls te Stomatorhinia Rondani

(JX)

Genus METALLEA van der Wulp (1880).

This genus has the centre of propleura haired, and the arista with very
short pubescence. The inner margins of the processes of fifth abdominal sternite
in males of the Australian species are furnished with closely placed short stubby
black bristles (Text-fig. 21), which are not so evident in the Oriental species
referred here, and the genital segment of the females has distinct spines as in
some other genera of the group.

Rhyncomyia differs from this genus in having the centre of propleura bare,
but in other respects is similar; the genus does not occur in Australia so far as
I am aware; the species referred here by Macquart possibly belong to Metallea.
I am of the opinion that the species described by Aldrich as illingworthi is
gracilipalpis Macquart, but an examination of the type of the iatter is essential
to determine if this supposition is correct. What the other species described as
Rhyncomyia dubia Macquart may be I do not hazard a guess.

In the following key I have used mostly colour characters, which though not
of outstanding significance as a rule, appear here to be reliable. My material is
not sufficient to permit of an extensive examination of the genitalia of the males,
the most dependable method for identification of the species.

Key to Species.

1. Frons of male at narrowest point about one-seventh of the head width; large spacies,
about 10 mm. in length, with a dark mark on cheek in front of the raised part;
parafacials with black setulose hairs; cheeks entirely yellow haired; fourth
visible abdominal tergite with two series of long bristles on disc ..........
eee aia fi G a. station le sugars CARR ore N eeu nT Motoke nevieneceiate; obey aie teel ener evens robusta Aldrich

Frons of male reduced to a mere line, at narrowest point not one-twelfth of the

ETS CUM VAL CUTTS tee ees ia eel 2 Lah, Seer PE SN Luan SU IEAM a traltet efisge vemeeueuanetey uel ceetal eats ese la jer opsieWone 2
Be IME acl Sati ee er IICER CREO In or nOreroier icin orton ofointcharcinibih, O-atoldl Gino, cecictals IG inmromica 3
CTIA CSPI pee a nec! cirnirs) (slice ov cuisie Topralt eule\ calmer Pare syslrepatenekanel sian avaheieacrcmemeusete ners aushorenateaicteteraton a 6

3. Entire parafacials and cheeks with fine yellow hairs, those on the former almost
imperceptible unless under a strong lens, the ground colour of face, parafacials,
and cheeks yellow; abdomen testaceous yellow, with a dark dorsocentral vitta or
series of spots, and dark spots on lateral curved portions of tergites, sometimes
with these markings much reduced, and at other times with them much diffused
aynl treme! Lrogeceob ooo Us ootdiccoddvacoguacbaadcodocDG illingworthi Aldrich

330 NOTES ON AUSTRALIAN DIPTERA, Xi,

Either the parafacials or cheeks, or both, partly or entirely black haired ...... 4

4. No black hairs on cheeks except directly below parafacials; abdomen testaceous
yellow, with a black dorsocentral vitta, a black transverse band close to apex

of third tergite, most of fourth black, and a black postspiracular spot on tergites

2 and 8, the incurved lateral portions of basal three tergites yellow ........
RRO DROS ERO TORO EPEC CR AT Gree GIGI AOR ERO OT CR AICIO NG rote) Gc oro Broa Oke rnaserd cin insularis, n. sp.

Most or allso£ the hairs onkcheeks) black Or fuSCOuSieaeieiee ae cieeie eto eine 5

5. Cheeks and parafacials entirely yellow; two or three of the abdominal tergites quite
extensively and conspicuously testaceous on lateral anterior—portions of disc

Sofas Raveleo'l elena eS Ea ae aS AGE ST CROAT OH MCN EW ETL Suis is oder ch le etude woigetPORMRE Ce Beps Mes te lees aerate nigribarba Aldrich

Cheek with a mark at middle, and the upper posterior part, fuscous, the parafacials
slightly darkened along eye on lower half .................. puncticeps, 1. sp.

6. Parafacials and cheeks yellow haired, sometimes with a feweof the parafacial hairs
danke remokamlan® eliymmeyiellOwarcvecislecicelsirrccsteehaionetenen nen terse illingwortnhi Aldrich
Parafacials with the hairs all black; femora fuscous ........................ a

7. Small species, not over 8 mm. in length; parafacial hairs fine .. nigribarba Aldrich?
Larger species, over 9 mm. in length; parafacial hairs strong .... insularis, n. sp.

: METALLEA ROBUSTA Aldrich.
Proc. U.S. Nat. Mus., 1926, lxix, art. 22, p. 9.

I have seen only the two specimens of this species from which the species
was originally described. The abdominal bristles are stronger than in any other
species and the frons of male is broader, as indicated in the above key.

Western Australia.

Type in United States National Museum.

METALLEA ILLINGWORTHI Aldrich. (Text-figures 21 and 22.)

Proc. U.S. Nat. Mus., 1926, lxix, art. 22, p. 7; ? gracilipalpis Macq., Mém. Soc.
Sci. Lille, 1854, p. 129, (1855); Dipt. Exot., Suppl. 5, p. 109, Pl. vi, fig. 3, 1855.

This is evidently a common species in eastern Australia as I have a long
series from New South Wales, Queensland, and South Australia. I am almost
eertain that this is gracilipalpis Macquart, but having received a name which we
are positive of in its application I use it pending an examination of the type
specimen of Macquart’s species if it is still available. Hypopygium as in
Text-fig. 22. Bezzi’s M. divisa is probably the same.

Text-figure 21. One process of fifth sternite of Metallea illingworthi.
Text-figure 22. Hypopygium of Metallea illingworthi. a, side view;
b, forceps from behind.

METALLEA INSULARIS, 0. Sp.

Male and female.—Head testaceous; occiput black, and with grey dusting
except below level of eyes; frontal orbits in male fuscous almost to bases of
antennae, and parafacials slightly darkened along eyes, the dark parts of both

BY J. R. MALLOCH. 331

grey dusted, cheek in same sex with a very faint dark central mark; orbits,
parafacials, and cheeks in female less noticebaly darkened than in male; antennae
brownish yellow; palpi yellow. Thorax metallic blackish green, with bronzy or
coppery reflections, and slight whitish dusting, the hairs and bristles on dorsum
inserted in dark dots, the dorsal vittae faint. Abdomen of male as described in
key, hypopygium black, with greenish tinge, entire surface with slight whitish
dusting; female with the abdomen more extensively blackened apically, the
hairs and bristles on dark parts in both sexes inserted in dark dots. Legs black,
femora with greenish tinge on parts, tibiae and bases of at least the fore and mid
tarsi reddish yellow. Wings greyish hyaline, yellowish at bases. Calyptrae and
halteres yellow.

Male.—F rons linear above, inner orbital series of bristles strong, outer parts
of orbits with black setulose hairs, similar hairs continued on parafacials to, or
almost to, level of vibrissae, the hairs on cheeks carried almost to vibrissae, black
anteriorly, yellow on raised part; numerous black setulae above vibrissae; arista
pubescent. Thorax with four bristles on presutural area, the usual weak anterior
sublateral bristle absent; dorsocentral bristles 2 + 4; one propleural and one
stigmatal bristle, both strong. As usual in the Australian species of this genus
there is no ventral bristle on the mid tibia in this sex; hind tibia with about
six irregular anterodorsal, one anteroventral, one posterior, and three postero-
dorsal bristles. Wings normal, one setula below and two or three above at base
of third vein.

Female.—Frons at vertex about one-fifth, at anterior margin over one-third,
of the head width, interfrontalia of uniform width on entire length, orbits much
widened anteriorly. Hind tibia with 4-5 anterodorsal, two posterodorsal and
two anteroventral bristles.

Length, 9-10 mm.

Type male, and allotype, Milson Island, N.S.W.

METALLEA NIGRIBARBA Aldrich.
Proc. U.S. Nat. Mus., 1926, |xix, art. 22, p. 10.

A female, which is evidently of this species, is from EHidsvold, Queensland
(Bancroft). I have examined the type male of this in the United States
National Museum. It is from eastern Australia.

METALLEA PUNCTICEPS, N. Sp.

Male.—A much darker species than any of the others, being without noticeable
testaceous markings on the abdomen. The markings on the head are constant in
the three males before me and appear to be of specific import. The abdomen is
blackish green, with whitish dusting through which the coppery reflections show
faintly on the sides of the tergites, the dorsum has a trace of a dark central vitta,
and there are dark dots at bases of the hairs and bristles only on the sides of the
tergites. In other respects similar to insularis.

In nearly every case is there a shorter bristle below the propleural one in the
type series of this species, while there is such a bristle on one side only in the
type of insularis and none on the allotype.

Length, 6-7:25 mm.

Type and two paratype males, Perth, Western Australia (Nicholson).

(0)

332 NOTES ON AUSTRALIAN DIPTERA, Xi,

Genus CHLORORHINIA Townsend (1917).

This genus is common in the Orient. The only species occurring in Australia

is readily distinguished from any other of the subfamily by its brilliant metallic
blue colour, besides the characters cited in the foregoing key.

CHLORORHINIA VIRIDIS Townsend.
Rec. Ind. Mus., 1917, xiii, p. 191.

The dorsum of the abdomen is largely opaque black in most specimens of the

species.
Localities: Illawarra and Sydney, N.S.W.; Kuranda and Cairns, Queensland.

Genus RHINIA Robineau-Desvoidy (1830).

This genus is very similar to Stomatorhinia in all particulars, but besides
the presence of only two bristles on the upper hind angle of mesopleura, which
is rarely departed from, instead of a complete series on its entire length, the males
have invariably a series of long bristles on the hind margin of the first visible
tergite, usually divided more or less at middle, which are not developed in the
other genera of the subfamily, and the hind tibia has a series of regular short
anterodorsal setulae with one or two slightly longer. In the species before me
there is a large glossy black undusted area covering the lower anterior angle of
mesopleura and the anterior portion of sternopleura in Rhinia, while in
Stomatorhinia the entire pleura is densely grey dusted. The presence of this
glossy area readily distinguishes the species listed below from those of the
species of Stomatorhinia occurring in Australia, but it may not hold throughout
the genus in other faunal regions.

RHINIA QUADRINOTATA Bigot.
Ann. Soc. Ent. France, 1874, p. 238.

I identify as this species several specimens amongst my Australian material.
These have been carefully compared with specimens from the Philippine Islands
and appear to agree perfectly with them, there being no structural or other
differences.

Localities: Cairns, N. Queensland (Dodd); Sydney, N.S.W.

Genus STOMATORHINIA Rondani.

This genus is most closely related to Rhinia, but is distinguished as stated
in the key and under the preceding genus.

The species of the genus are difficult to separate satisfactorily and even
the two Australian species require further elucidation, though the two forms
accepted in this paper are undoubtedly distinct, the species listed as subapicalis
being very similar to one or two others occurring in the Orient.

I present a synopsis of the species below.

A. Testaceous yellow species, with yellow palpi, orbits and lower half of face
castaneous, thorax largely fuscous, obscured by dense whitish grey dusting,
scutellum testaceous, with a conspicuous black spot at base of each of
the four marginal bristles; no well developed posteroventral bristles on
basal half of hind femur of female; wing without an evident preapical dark
costal mark; black spots of frontal orbits biseriate ............ pallida, n. sp.

BY J. R. MALLOCH. 333

AA. Greenish black species, with black or dark brown palpi, anterior half of cheeks
as well as lower half of face glossy black or castaneous, the scutellum at
most testaceous on margin, and with inconspicuous dark dots at bases of the
four marginal bristles; abdomen with lateral testaceous marks on basal two
or three tergites; hind femur with one or two long bristles on basal half of
posteroventral surface; black setigerous spots on frontal orbits triseriate
ATU CTO Wi Vane Meeytoyayevailearaiej os teisaitorrer ey elfails oyeliousdehele #is)susvere.s vee (euexenciiey's subapicalis Macquart

STOMATORHINIA PALLIDA, 0. SD.

Female.—Head testaceous yellow; orbits castaneous, densely whitish dusted.
with two series of glossy black piliferous spots; face glossy, with a broad central
band of whitish dust, the lower undusted half castaneous; parafacial white dusted,
with a few piliferous glossy spots and, near lower extremity, a large glossy
mark; cheeks densely white dusted, anterior half with an oblique glossy area;
antennae testaceous, third segment more or less browned; palpi testaceous. Thorax
largely fuscous, densely whitish grey dusted, the entire pleura covered with
white dust, and, like the dorsum, with many dark piliferous spots; scutellum
testaceous, dusted and dotted like the mesonotum, and with a conspicuous black
spot at base of each of the four marginal bristles. Abdomen testaceous, with dark
apices to the tergites, dusted and dotted like thorax, the dots at apices of tergites
most evident. Legs testaceous, apices of tarsi darkened. Wings hyaline. Calyptrae
white. Halteres yellow.

Structurally similar to the other species of the genus, but in the two
females I have seen there are no noticeable bristles at base of the posteroventral
surface of hind femur.

Length, 6 mm.

Type, Cairns, N. Queensland. One paratype, same locality (J. F. Illingworth)
[U.S.N.M.].

STOMATORHINIA SUBAPICALIS (Macquart). (Text-figure 23.)

Mém. Soc. Sci. Lille, 1846, p. 98 (1847); Dipt. Hxot., suppl. 2, p. 82, pl. v,
fig. 4, 1847; australis Walker, Cat. Dipt. Brit. Mus. iv, p. 809, 1849; murina Schiner,
Reise Novara, Dipt., p. 309, 1868.

This species resembles lunata Fabricius, but the dorsum of thorax has many
black piliferous dots and no evident vittae, the pleura are more densely grey

Text-figure 23. Hypopygium of Stomatorhinia subapicalis;
a, side view; b, forceps from behind.

dusted and more conspicuously black dotted, the wing has a more or less evident
dark preapical costal cloud, and the mid femur has a series of short stout spines
at apex on posteroventral surface which is not present in the African male before
me. Hypopygium as Text-fig. 23.

Length, 6-8 mm.

334 NOTES ON AUSTRALIAN DIPTERA, Xi,

Localities: HEidsvold, Queensland; Sydney and Botany Bay, N.S.W.; Beacons-
field, Victoria and Myponga, S. Australia.

STOMATORHINIA CRIBELLATA Bezzi.

Stomorhina cribellata Bezzi, Bull. Ent. Res., 1927, xvii, p. 233.
I have not seen this species, unless it may be the species I accept as
subapicalis. It is not pallida.

STOMATORHINIA DISCOLOR Fabr.
Ent. Syst., iv, p. 320, 1794. (Doubtful record.)

STOMATORHINIA XANTHOGASTER Wied.
Aussereurop. Zweifl. Ins. ii, p. 349, 1830. (Doubtful.)

Subfamily Sarcophaginae.

The members of this subfamily occurring in Australia have been competently
handled by other workers so that it is unnecessary for me to do more than mention
some unconsidered characters here.

In no work dealing with the genus Sarcophaga has there been any mention
made of such characters as the hairs on prosternum, centre of propleura, and
centre of the declivous portion of thorax above calyptrae. I find that by the use
of these characters, which are present or absent in certain groups, it is possible
to associate more readily the sexes than it is by any other means, and believe
that the adoption of their use in classification would simplify the identification of
the species, at least in so far as the females are concerned.

I have on a previous page dealt with the structure of the hypopygial forceps
as an indication of group location. It is also possible to use the structure of the
genital segments of the females as specific indices, but very little has been done
in this respect by any worker in this genus. There are some very characteristic
features of these segments in some of the species, possibly arising from adaptations
to some peculiarity in oviposition, but whether they lend themselves to grouping
of the species as well as to identification of individual species I have not
determined as yet.

Subfamily Metopiinae.

This subfamily has most frequently been placed in the Tachinidae, but it
undoubtedly has closer affinities with Sarcophaginae than with any subfamily
of Tachinidae.

The second abdominal sternite is fully exposed and overlies the lateral margins
of the second tergite as in other subfamilies of Calliphoridae, but the arista is always
bare or pubescent, the lower calypter is always bare on upper disc and in most,
if not all, genera bulged up near base, with an inflated appearance; the sterno-
pleurals are frequently 1 + 2 or 3; and the notopleurals are two in number. The
small knob-like protuberance before base of wing on upper part of pleura is well
developed and stands out from the side more than does the upper margin of the
pleura, as in Calliphoridae, while in Tachinidae it is as a rule Jess developed and
about level with the upper edge of pleura.

The species so far as known occur in the immature stages in nests of solitary
bees, but the life-histories are but little known, and whether they are parasites
or scavengers is not definitely known.

BY J. R. MALLOCH. 335

There appear to be two genera in my Australian material, both of them
already known to science. They may be distinguished as below.

A. Vibrissae not differentiated, the bristles of uniform length from some distance below

vibrissal angle to well above it ....... Nibeiencueesiuuabereneeevacenets Miltogramma Meigen
AA. Vibrissae well developed and quite distinct from the setulae above and below
ELVGTYMMMR PAR Mone Ve etecsids i) overs: oicis ohenseemensteteneheonsrone tal'ensis te Protomiltogramma Townsend

Genus MinToGRAMMA Meigen.
I appear to have two species of this genus, but represented by single specimens.

Genus PROTOMILTOGRAMMA Townsend.

I have seen the genotype of this species in the United States National
Museum, and have several specimens which are either the same or a closely
allied species. Pending receipt of more material I leave these genera in abeyance.
The genotype is cincta Townsend, described from Queensland.

Unrecognized Species.

Calliphora elliptica Macq. Mem. Soc. Sci. Lille, 1846, p. 99 (1847); Dipt. Exrot.,
suppl. 2, p. 83, pl. v, fig. 6, 1847.

Pollenia ruficornis Macq. Mem. Soc. Sci. Lille, 1846, p. 101 (1847); Dipt. Hxot.,
suppl. 2, p. 85, pl. v, fig. 8, 1847.

Pollenia viridiventris Macq. Mem. Soc. Sci. Lille, 1846, p. 100 (1847); Dipt. Ezxot.,
suppl. 2, p. 84, pl. v, fig. 9, 1847.

Pollenia tasmanensis Macq. Mem. Soc. Sci. Lille, 1850, p. 227 (1851); Dipt. EHzot.,
suppl. 4, p. 254.

Pollenia ruficornis Macq. Mem. Soc. Sci. Lille, 1850, p. 227 (1851); Dipt. Exot.,
suppl. 4, p. 254.

Pollenia moretonensis Macq. Mem. Soc. Sci. Lille, 1854, p. 186 (1855); Dipt. Exot.,
suppl. 5, p. 116, 1855.

Anastellorhina bicolor Bigot. Bull. Hnt. Soc. France, 1885, p. xxvi.

Rhyncomyia incisuralis Macq. Mem. Soc. Sci. Lille, 1850, p. 214 (1851); Dipt.
Exot., suppl. 4, p. 241, pl. xxii, fig. 8.

Rhyncomyia dubia Macq. Mem. Soc. Sci. Lille, 1854, p. 129 (1855); Dipt. Hzot.,
suppl. 5, p. 109, pl. vi, fig. 4, 1855.

Rhyncomyia tigrina Bigot. Ann. Soc. Ent. France, 1874, Ser. 5, iv, p. 242.

Principal recent papers on Australian Calliphoridae as limited in catalogue.

ALDRICH, J. M.—Proc. U.S. Nat. Mus., 1925, Ixiv, art. 18. Deals with Wicrocalliphora.
Id., Proc. U.S. Nat. Mus., 1926, lxix, art. 22. Deals with Metallea.

Buzzi, M.—Bull. Ent. Res., 1927, xvii, pt. 3. Keys to species of South Pacific Islands
and Australia, with a few descriptions of new Australian species.

Harpy, G. H.—Proc. Roy. Soc. Qld., 1925, xxxvii (1926). Deals with the Australian
species of Calliphora.

JOHNSTON, T. H., and Harpy, G. H.—Proc. Roy. Soc. Qld., 1922, xxxiv, No. 3. Biology.
Id., 1922, No. 10. Synonymic list of species.

MALLocH, J. R—Ann. Mag. Nat. Hist., 1926, (9), xvii, p. 507.

Patton, W. S.—Phil. Jour. Sci., 1925, xxvii, No. 3. Deals with Calliphora, Lucilia and

' Chrysomyia sens. lat.

NOTES ON AUSTRALIAN DIPTERA. No. xii.

By J. R. MAtLtocu.
(Communicated by Dr. I. M. Mackerras.)

(Two Text-figures. )
[Read 3lst August, 1927.]

In this part of the series of papers which I have been intermittently presenting
I include some data upon the family Tachinidae which may be of interest to
students of the group who intend to take up a more intensive survey of these
interesting and, in some cases at least, valuable economic species.

The group appears to be a comparatively recent one, and, as in most such
groups, the generic and specific limits are not as sharply drawn as in the
older groups, and there is a greater diversity of characters, with a corresponding
lack of distinctness in some of these, than is present in more definitely established
groups and species. It is to be regretted that much of the work already done
has been based upon rather trivial and variable characters, and, in addition
to that, practically all of it done outside of Europe has been grafted upon the
basis supplied by earlier work upon the fauna of that continent, and without
considering either the fauna of the world as a whole or that of each particular
faunal region as a separate unit. Consequently many of the species described
as belonging to particular genera do not belong to, and are often quite distinct
from, the genera they were originally placed in, having been relegated to their
generic position without a proper consideration of many important characters,
because of a lack of knowledge of the groups and their characters. Even the
major groups which have in many cases been accepted as of family and subfamily
status are not entitled in most cases to such distinction, and herein I include
in the family the group Dexiidae of authors, and such concepts as Rutiliinae, etc.,
limiting the family as follows: Hypopleura with distinct bristles in one or more
series below the spiracle, or adjacent to it; postscutellum quite prominently
convexly developed immediately below the scutellum; lower calypter always
much larger than upper; none of the abdominal sternites overlapping sides of
tergites on their entire length. Usually the small knob-like protuberance
immediately in front of and below base of wing is inconspicuous, but in some of
the groups it is as prominent as in Muscidae.

There are a number of groups which may be accepted as subfamilies or tribes,
such as Dexiinae, Rutiliini, Phasiinae, etc., and hereinafter I deal with one
segregate which has been accorded tribal rank by Townsend and the present
writer, and which is essentially Oriental and Australasian so far as I am at
present aware.

The figures which I presented in my recent paper (These ProcrkEpiInes, 1927)
on the Australian Calliphoridae will be found useful in the study of this
family also, most of the characters shown therein being used here also.

BY J. R. MALLOCH. 337

Tribe Palpostomini.

This group is distinguished from its allies by the following combination of
characters: Scutellum with four strong bristles; prosternum setulose; lower
calypter narrow, rounded at apex, and standing well clear of lateral basal angle
of scutellum, its inner basal portion straight; third wing vein ending in costa
close to apex of wing; first wing vein bare; central portion of upper occiput
with two groups of short black setulae on lower half or more; palpi well developed.
The proboscis may or may not have the apical labellae separated and palp-like.

There are three genera at present known to me, two of them occurring in
Australia. I give below a key for their recognition.

Key to the Genera.

1. Centre of propleura with setulose hairs ......................- Hustacomyia, n. gen.
GentrerOLspropleunTar DATE) ec se ce per caemen ete sitreitatel avereraioue chellltene tetera euetshioueiete terezene 2

2. First posterior cell of wing usually closed and distinctly pedunculate, rarely closed
AtexXtreme Mancino wm seein oeces cele cichee Palpostoma Robineau-Desvoidy

First posterior cell of wing distinctly open in margin of wing ................:.
6) a JOE CREE Cea CRORE oral oR Rear ie i OnE eS Te grhL en ical Fiaot ane AU ne Ura Ochromeigenia Townsend

Genus OCHROMEIGENIA Townsend.

There is but one described species of this genus, ormoioides Townsend, which
occurs in the Orient and is parasitic upon beetles.

Genus EHUSTACOMYIA, n. gen.

This genus is very similar to Ochromeigenia, having the first posterior cell
of wing open, but it differs from both the other genera in having the central
part of the propleura with numerous setulose hairs, and the setulae on base of
third vein confined to the node both above and below, instead of only below. In
addition to these characters the parafacial is wider, being noticeably wider
below than is the third antennal segment, and the arista is much shorter, being
barely as long as antennae, and more thickened on basal half, with almost
indistinguishable pubescence. Genotype, Hustacomyia breviseta Malloch. Named
in honour of the late Dr. Eustace W. Ferguson, who submitted the material.

EUSTACOMYIA BREVISETA, nN. Sp.

Male.—Head yellowish ochreous, occiput, ocellar spot, frontal orbits, and
parafacials, fuscous, with grey dusting, third antennal segment and arista
fuscous, palpi ochreous. Thorax fuscous, more yellowish on humeral angles
and apex of scutellum, the surface with changeable grey dusting, mesonotum with
four linear black vittae, which are interrupted at suture and cease behind middle,
and a blackish mark laterad of these on each side at suture, all of which are best
seen when the thorax is viewed from behind; metanotum greyish dusted. Abdomen
ochre-brown, with a darker central vitta, and dots at bases of the hairs and
bristles, the anterior and posterior margins of tergites pale grey dusted. Legs
ochreous yellow, fore and hind femora, and apices of tarsi, infuscated. Wings
slightly smoky. Calyptrae pale brownish. Halteres dusky yellow.

Frons at anterior ocellus fully as wide as third antennal segment and about
half as wide as height of cheek, vertical and ocellar bristles very short, orbits
with a series of short bristles along inner margins, and short setulose hairs
laterad of these which are continued downward over parafacial in about six
series, the parafacial at middle about 1:5 as wide as third antennal segment,

338 NOTES ON AUSTRALIAN DIPTERA, Xii,

the latter fully twice as long as second segment; arista thickened to about
middle, almost bare; cheek about one-third as high as eye; vibrissae short, with
numerous black bristles adjacent to them; proboscis without differentiated apical
processes. Thoracic dorsum with the hairs longer and stronger than in Palpostoma,
the presutural acrostichals as long as the dorsocentrals, in other respects as in
that genus. Tibial bristles rather weak. First posterior cell narrowly open.

Length, 6 mm.

Type, Sydney, N.S.W., December 16, 1923.

Genus PALPpostoMA Robineau-Desvoidy.

This genus is known only from Australia and Tasmania, and was originally
placed in the heterogeneous assemblage “Muscidae Testacea’” by its describer,
but has little in common with most of the other genera therein included, these
being mostly Calliphoridae, the only character they had in common being their
testaceous yellow colour.

Aldrich in 1922 dealt with the genus, briefly redescribed the two known
species, and added one new one.*

I have examined his material and, having a number of additional specimens
as well as some new data upon the genus, I present some notes herewith.

In redefining the genus Aldrich gave a comprehensive description of the
thoracic bristling, which unfortunately does not apply in all respects to the
material I have examined, there being in at least one species more than two
prosternal bristles, in nearly all specimens there are two intra-alars, and the
posterior mesopleural bristles are more than four in number. It is also note-
worthy that the palp-like apical labellae of the proboscis are not always
conspicuous, there being specimens in which they are even undivided, or
almost so.

It is extremely difficult to decide from the material now in my hands exactly
how many species are represented, as some are from widely separated localities
from those that yielded others, and there is but one sex represented from certain
of those localities. No taxonomist is more ready to admit than I am the dangers
attendant upon the elucidation of species from a small amount of material, and
without a knowledge of the habits, or biology, of a species, or the circumstances
of its occurrence in nature, and were I to attempt the presentation of complete
data upon this material I fully realize that there would be considerable risk of
falling into error in my conclusions as to specific distinctions, so I refrain from
making a complete survey pending receipt of further accessions.

One of the specimens before me appears to represent a species quite distinct
from any of the three in Aldrich’s material and is described below. In addition
to it there are apparently four other species in my possession, one of these is
represented by 10 specimens, all taken in Sydney, N.S.W., and appears to be
distinct from the form accepted by Aldrich as testacea Robineau-Desvoidy, though
it seems to be even more likely to prove the true testacea than the other one.
Two females from Sydney belong either to two distinct species, or if they belong
to but one they present an unusual dimorphism, one having the frons very
little wider at anterior ocellus than the distance across the posterior ocelli and
not as wide as third antennal segment, while the other has it twice as wide as
either of these standards. The genus appears to me to present an interesting
field for study by some person having access to the insects in their natural

-* Proc. U.S. Nat. Mus., 1922, 1xii, art. 11, pp. 2-5.

BY J. R. MALLOCH. 339

surroundings; any decisions based upon material such as I have, and under
laboratory conditions, must of necessity be largely conjectural.

In a paper published recently in the Philippine Journal of Science, Townsend
has erected the genus Pseudopalpostoma for the reception of Palpostoma desvoidyi
Aldrich. This genus is absolutely uncalled for, as none of the characters cited
for its distinction are absolute, and some of them, such as the single strong
median marginal bristle on the second apparent abdominal tergite, are abnormal.
Bvery student who has a desire to keep systematic entomology on a practical
basis must deprecate this erection of illusory genera, and especially in faunal
regions which are terra incognita to the worker who merely dabbles in inventions,
without aiding either beginners or those who are really versed in the taxonomy
of the group. The literature of this family is in worse condition than that of
any other, and unless one is prepared to disregard most of it and proceed with
his work along conservative lines he must perforce devote a tremendous percentage
of his time to untangling the figments of imagination tendered as serious
contributions to our knowledge of the family since the latter part of the last
century.

PALPOSTOMA APICALIS, N. Sp.

Female.—Shining fulvous-testaceous. Ocellar spot and a streak on each side
of occiput above blackish; frontal orbits and upper part of parafacials slightly
infuscated, and, like the face, white dusted; third antennal segment brown except
at base. Thorax without evident dorsal dusting. Abdomen fuscous and with
rather dense brownish dusting on apical three tergites, the basal tergite entirely
fulvous-testaceous. Apices of mid and hind femora blackened above, the mid
pair least distinctly so and to a lesser extent. Wings hyaline. Calyptrae and
halteres yellow.

Frons at vertex about one-fifth of the head width, about twice as wide as
third antennal segment, and three times as wide as distance across posterior
ocelli; the forwardly directed upper orbital bristles and the ocellars very weak,
the four verticals well developed; third antennal segment over three times as
long as second and not as wide as half the height of cheek; arista subnude;
cheek fully one-third as high as eye. Thoracic dorsum more sparsely haired
than usual, the bristles fine but distinct, anterior intra-alar short. Legs slenderer
than usual, the tibial bristles all short and weak, none on anterodorsal surface
of fore pair. No evident bristles at apex of fourth abdominal tergite. First
posterior cell closed in margin of wing, not petiolate.

Length, 5-5 mm.

Type, Woy Woy, N.S.W., September 22, 1923 (Mackerras).

A rather slender and weakly bristled species, distinguished from any known
to me by the dark apices to the mid and hind femora. I hope to obtain enough
material to deal further with the genus at some future time.

I have been unable to decide just what group to place the following genus in,
but the evident though short hairs on the arista would apparently indicate that
it is dexiine in its affinities. There are several Oriental species allied to the
one now dealt with, but these cannot be completely described here.

Genus SEMISUTURIA, Nn. gen.
Generic characters.—I{f one considers the very short haired arista as
indicating group affinities, this genus runs to Morphomyia Rondani in Williston’s
Q

340 NOTES ON AUSTRALIAN DIPTERA, Xii,

key to the genera of Dexiinae of North America. A comparison with the
genotype, tachinoides Fallen, shows many points of resemblance, but the abdominal
tergites in the new genus are not separated by distinct sutures except laterally,
while in Morphomyia and other genera known to me the sutures dividing the
tergites are complete. If, on the other hand, one disregards the short aristal
hairing as of no value in indicating group relationships, and assigns the genus to
Tachininae, the genus will run to Oestrophasia in the same manual, but here
the same character as above indicated will suffice for its recognition. A summary
of the characters of the new genus is as follows: Arista very short haired; eyes
with sparse microscopic hairs; face without a vertical carina; antennae in
profile inserted close to middle of eyes; cheek not one-third of the eye height;
vibrissae situated a little above mouth margin, some bristles above them;
parafacial bare; proboscis short; palpi well developed. Thorax with 2 or 3 + 3
dorsocentral bristles, 1 or 2 + 2 acrostichals; only two bristles on presutural area,
the posterior sublateral lacking; postsutural intra-alars two pairs; prosternum,
propleura, postalar declivity, and metanotum, bare; sternopleurals 1 + 1; posterior
spiracle with pronounced posterior flap. Abdomen ovate, or elongate ovate, the
tergites apparently fused centrally; a pair of macrochaetae on middle of hind
Inargin of each tergite except on first in two species. Costal thorn present; inner
crossvein well before apex of first vein; bend of fourth vein subangular; first
posterior cell open, its apex quite close to wing tip; third vein setulose at base;
other veins bare; lower calypter lobulate at base on inner margin and lying
close to scutellum. Genotype, Semisuturia australis, n. sp.

SEMISUTURIA AUSTRALIS, n. sp. (Text-figure 1.)
Female.—Fulvous testaceous, shining, third antennal segment and aristae
black. Wings yellowish hyaline.

ee seael Die ea

Text-fig. 1. Apex of wing of Semisuturia australis.
Text-fig. 2. Wing of Amenia imperialis, male.

Frons at vertex a little less than one-third of the head width, gradually
widened to anterior margin where it is about two-fifths of the head width,
orbits shining, whitish dusted like the parafacials, upper bristle sloping backward,
two forwardly directed bristles on upper half, and about six inwardly curved
bristles along inner margin of each orbit; ocellar bristles over twice as long as
postvertical pair, divergent and forwardly directed; eye sparsely haired; cheek
over one-third of the eye height, the raised part with three or four strong
bristles above; several short bristles above vibrissa; arista pubescent; palpi of
moderate length. Thorax with 2 + 3 dorsocentrals and 2 + 2 or 3 acrostichals,
three humerals, two bristles on presutural area, two pairs of intra-alars, and no
bristle between the posterior dorsocentral and postalar bristles; scutellum with
6-8 marginal bristles. First tergite without central apical bristles, second with a

BY J. R. MALLOCH. 341

strong pair, third, fourth, and fifth, with practically complete series of apical
bristles, and none on disc. Amnterodorsal series of bristles on fore tibia quite
pronounced; ventral bristle on mid tibia very short and weak; hind femur with
one or two bristles before and beyond middle on anteroventral surface. Inner
crossvein well before apex of first vein and close to middle of discal cell; apical
venation as in Text-fig. 1.

Length, 6-5 mm.

Type, Hidsvold, Queensland, December, 1922.

There are several Oriental species of this genus known to me, none of which
I have been able to identify with published descriptions. These will be dealt
with in another paper published in England. To establish these species, some
of which may occur in Australia, I. present below a key to the whole of the
species of the genus now before me, reserving the full descriptions for the other
paper.

Key to the Species.

1. Bend of fourth wing vein with a quite conspicuous vein emanating from it at the
angles which 1Sh Quitem Sharpie acest crea custeie enol cveverel siecle een steven veisiiel el cveroneneueia rete 2

Bend of fourth wing vein without any evidence of a vein emanating from it .... 4

2. Abdominal tergites except first each with a pair of discal macrochaetae in line
with the apical central pair; metanotum and abdomen glossy black, basal, and

sidesof, second: tergites yellows eon sienieccleromlcnens sic sieuerelcleienel nitidiventris, n. sp.
Abdominal tergites except fourth without discal macrochaetae; abdomen not coloured
ENS) EWSKONE) Moiplom cin aor a 4016 CIOL Late Oe cote Go OBIa ce Onin Did pb oomd Do ddoo GOGO OOK 3

8. Third antennal segment black; abdomen glossy brownish black, apex of fourth
Werte AKMIONY CosdaoosdodseooosdocosdiogououabuoououMaodG OOD nigricornis, Nn. sp.
Antennae entirely yellow; abdomen yellow, brownish apically .... flavicornis, n. sp.

Ame Thirdeantennaluseeim erica a cko aeraciensinciciensieciele icten-ten ete wonalel ie -ikeleheNeteiel ol cuehetcloMel hek-ioiel shale 5
Third antennal segment not black, yellow, or at most brownish ................ 216

5. Entire insect except third antennal segment and aristae fulvous testaceous ......
PR afc etree EMT ieee BCs Aral Wome tioe aicgt NI alist iets niepere Malick’ aepsasintetsncnaiscened ste australis, n. sp.
Thorax with black dorsal vittae, abdomen glossy black on dorsum, apex of fourth

Witsoe Wests) cAI |S oiosocccuodndDadoubODod boo go GadcaoD pahangensis, n. sp.
6. First visible tergite with a pair of macrochaetae on middle of hind margin; outer
erossvein, and costa apically, without clouding .............. hyalipennis, n. sp.

First visible tergite without developed macrochaetae on middle of apical margin;
outer crossvein, and costa beyond apex of second vein, clouded ............
fk ive) Ur OR nea STE EM ag eR ea oe On AR TS st Pe oi ORO rue triangulifera, n. sp.

I give below the type localities of the new species and the collections in which

they are to be found.

SEMISUTURIA NITIDIVENTRIS, TD. SD.

Type, Lubok Tamang, Pahang, Federated Malay States, 3,500 feet, June 9,
1923 (H. M. Pendlebury), British Museum.

SEMISUTURIA NIGRICORNIS, N. Sp.
Type, Gunong Tahan Padang, Pahang, Federated Malay States, 5,550 feet,
January 3, 1923 (H. M. Pendlebury). British Museum.

SEMISUTURIA FLAVICORNIS, N. Sp.
Type, Singapore, Straits Settlements (C. F. Baker). Author’s collection.

SEMISUTURIA PAHANGENSIS, Nn. Sp.
Type, Gunong Tahan, Pahang, Federated Malay States, 6,000-7,000 feet,
December 13, 1921 (H. M. Pendlebury). British Museum.

342 NOTES ON AUSTRALIAN DIPTERA, Xii,

SEMISUTURIA HYALIPENNIS, 1. SD.
Type, Singapore, Straits Settlements (C. F. Baker). Author’s collection.

SEMISUTURIA TRIANGULIFEBA, Nl. SD.
Type, allotype, and two paratypes, Cuernos Mts., Negros, Philippine Islands
(C. F. Baker). Author’s collection.
I have another specimen which appears to belong to a distinct species, from
the Federated Malay States, but it is teneral and I do not care to describe it.

Tribe Ameniini.

This group is a difficult one to place satisfactorily. Most authors have placed
it in the same group as Rutilia Robineau-Desvoidy, but it does not appear to
me to be very closely related to that genus. In fact there is a very noticeable
difference in the structure of the postscutellum in the two genera I refer here
from that of typical tachinids, which Rutilia more closely resembles. In Ameniini
there is a very much smaller postscutellum than usual in Tachinidae, and, while
in the latter group there is always a rounded chitinized portion which bends
forward at upper margin, in the group now under consideration the chitin does
not noticeably bend forward, and the convexity is therefore much less pronounced.
Were the postscutellum totally lacking I might be inclined to refer the genera
involved to Calliphoridae, but in the latter the second abdominal sternite always
overlaps the lateral margins of the second tergite, while in Amenia and Stilbomyia
it does not. It is noteworthy that the small knob-like process in front of wing
base on the pteropleural suture is quite prominent and more like that of a
calliphorid than that of a tachinid, the typical forms of the latter group having
this knob about on a level with the one immediately below the notopleural suture.
A point of distinction between Ameniini and Calliphoridae is that the anterior
one of the two outermost bristles on the presutural area is in line with, or mesiad
of, the posterior one; in Calliphoridae it is as a rule laterad of it.

A definite conclusion as to the exact location of the Ameniini will depend
upon a knowledge of all stages, and pending this information I retain the tribe
in the Tachinidae.

Genus AMENIA Robineau-Desvoidy.

This genus contains species of a bright metallic blue or blue-green colour,
with conspicuous spots of white dust or tomentum on the thorax and abdomen.
The parafacials are bare, the prosternum, centre of propleura, mesonotum below
calyptrae, and the hind coxae above bases of femora, are furnished with fine hairs.
The centre of the declivous portion of thorax between base of wing and scutellum
is furnished with stiff hairs, the suprasquamal ridge is bare, and there are usually
four bristles on the presutural area, the anterior one of the posthumerals being
almost in line with or mesiad of the posterior one.

Dr. Engel in a recent paper recognized three species as occurring in Australia
and gave a diagnosis of their characters, but I do not agree with his conclusions
as to the species names. The species he calls leonina Fabricius is quite evidently
imperialis Robineau-Desvoidy. Wiedemann had before him the type specimen of
leonina when he re-described the species, and his very full and lucid description
does not apply at all to the species designated by this name in Engel’s paper, but
it does apply to that referred to as stictica Schiner in litt. It is quite clear from
an examination of Schiner’s description of parva and his remarks on leonina that

BY J. R. MALLOCH. 343

he misidentified the latter, placing imperialis and it under the same specific name,
and Engel apparently followed suit. I have attempted in this paper to rectify
what I consider to be errors of preceding workers, and at the same time separate
material before me on the basis of what I take to be specific characters. I have
what appear to me to be four species in the material available to me, all of it
from the United States National Museum collection, kindly made accessible by
Dr. Aldrich of that institution. In the determination of parva Schiner there is
some doubt, but it appears highly probable that the single specimen which I refer
here is correctly named, at least it is the only one which I can reliably distinguish
from leonina, except the forms without the dorsal thoracic vittae.

Key to the Species.

1. Thorax without evident submedian white dusted dorsal vittae in front of suture;
abdomen without trace of white submedian spots on second tergite, only the
one on each lateral curve present; frons of male at narrowest point much more
than twice as wide as distance across posterior ocelli; presutural acrostichals not

noticeably differentiated from the surrounding hairs ......................- 2
Thorax with a pair of very evident white dusted submedian vittae in front of
SHUN GBD 7S 8 ae iry cS IPR Bio TEs Osuna fer Seer ass -6.d ACA OL OkeeCr Otcl cent AR eRe Lat EG Tas DESIG MOTO cas eEtoTG Gece oad 3

2. Ocellar bristles well developed and widely divergent in male, the frons at vertex
in same sex five or six times as wide as distance across posterior ocelli ......
5'A50-0/ 850 NLA ORO ERR RRIGNC TE otOonunercher occa crural oorcic ore GIES etcioneee imperialis Robineau-Desvoidy

Ocellar bristles almost indistinguishable in male, the frons at vertex in same sex not
four times as wide as distance across posterior ocelli .......... dubitalis, n. sp.

3. Anterior postsutural dorsocentral bristles of thorax short and weak; postocular orbits
very little paler on upper half than below; second abdominal tergite with a pair
of submedian whitish dusted spots, sometimes faint .......... leonina Fabricius

Anterior postsutural dorsocentral bristles of thorax well developed and quite strong;
postocular orbits silvery white on upper half, contrasting with the yellow dusting
of the lower half; second abdominal tergite without submedian white dusted
spots, only the usual one at each lateral curve present .......... parva Schiner

AMENIA IMPERIALIS Robineau-Desvoidy. (Text-figure 2.)

This species is bright metallic green in colour, with the usual three white
spots along each side of thoracic dorsum, and there is a very faint dusting on the
anterior portion of thoracic dorsum which, however, is not in the form of vittae
and is visible only when seen from behind. There is no submedian pair of white
spots on the second abdominal tergite, only the lateral pair being present. The
fine hairs on the orbits laterad of the series of fine bristles are yellow in colour.

Structurally this species differs from the next one in having the frons wider,
and the submarginal cell of wing more obviously widened at middle than is the
case with that species (Text-fig. 2).

Length, 12-13 mm.

Localities: Cairns, N. Queensland (Illingworth), and Buderim Mt., Queensland.

AMENIA DUBITALIS, 0. SD.

This species is similar in colour to the above one but is smaller, and differs
as indicated in the key to species, and in the above notes on imperialis.

It is possible that with a large series of specimens available the distinction
will fail and the specimen may really represent only an aberration of imperialis.

Length, 10-5 mm.

Type, National Park, Sydney, N.S.W., November 1, 1902 (W. W. Froggatt).

344 NOTES ON AUSTRALIAN DIPTERA, Xii,

AMENIA LEONINA (Fabricius).

This species is apparently the commonest one of the genus and is readily
distinguished by the characters mentioned in the above key. The fourth visible
tergite is usually much darker than the others in both sexes which is not the
case in any of the other species.

The frons in the male is reduced to a mere line above, and that of the female
is about one-fourth of the head width at vertex, becoming wider anteriorly and at
the antennae about one-third of the head width, with the interfrontalia always
wider than either orbit, and the forwardly directed bristles quite weak.

Length, 9-12 mm.

Localities: Buderim Mt., Qld.; Kuranda, Qld.; Hamilton, Qld.; and Shoalhaven,
N.S.W.

AMENIA PARVA Schiner.

The specimen before me to which I assign this name is bright metallic blue-
green in colour, and readily distinguished from typical specimens of Jleonina,
which are normally deep blue or violet-blue in colour. The white upper portion
of the postocular orbits and lack of submedian white spots on second tergite as
well as the green colour of the fourth visible tergite, are the outstanding features
in the differences in the colour markings between this species and leonina, but
the second antennal segment is largely fuscous in parva and entirely yellow in
leonina, and while the fuscous cloud in the costal cell of wing ceases quite
abruptly a little beyond the humeral crossvein in leonina it gradually tapers off
apically in parva, the entire cell being evidently darkened. The fourth visible
tergite is the same colour as the others in parva. It is also noteworthy that all
the hairs on the head in parva are black, while in Jeonina those of the cheeks
and occiput are golden yellow. The interfrontalia of female is not wider than
either orbit.

Length, 9-5 mm.

Locality: Buderim Mt., Qld.

I give the lengths of the species in accordance with the material before me.
It appears pertinent to indicate that Schiner gives the length of parva as 2-2-5 lines,
which is about 4:25-5 mm., and Engel gives the length of this species as 8-11 mm.
As Engel had the type specimens of parva before him I accept his statement as
correct. It would appear that this genus would furnish an interesting field for
investigation to some resident entomologist just as all of the others would, and it
may develop that there are more species in Australia than is as yet suspected.

Genus STILBOMyIA Macquart.

This genus is very similar in almost every respect to Amenia, and the
character cited for their separation by Engel does not hold good for all the
species. This character is the length of the third antennal segment as compared
with the horizontal diameter of the eye. In Stilbomyia the third antennal segment
is given as greater than the horizontal diameter of the eye, while in Amenia it is
stated to be not over three-fourths of that diameter. I find, however, that in a
Chinese species of the former the antenna is about as short as in Amenia, and in
an Australian species the third antennal segment is about equal to the diameter
of the eye. The height of the cheek in Amenia is always much higher than the
entire length of antenna, whereas in Stilbomyia it is not as high except in the
Chinese species already referred to where the two are nearly equal. The frons

BY J. R. MALLOCH. 345

of the males of Amenia is very much narrower than in the females, and never
has strong forwardly directed bristles on the upper half on the outside, while in the
males of Stilbomyia the frons of the male is as wide as it is in the female, and in
both sexes there are at least two strong forwardly directed outer bristles on the
upper half; the females of Amenia have such forwardly directed bristles but they
are much weaker than in Stilbomyia. In both genera the abdominal sternites
are widely visible between the tergites apically, the exposed surface narrowing
considerably from apex to base.

Engel does not record this genus from Australia though he gives one species
as occurring in New Guinea. I have before me one specimen belonging to the
United States National Museum which is referable here.

STILBOMYIA OPULENTA (Walker).

This species may be the same as costalis (Walker), placed in the genus by
Engel, but the face is only slightly browned in the antennal fovea, not dark brown,
and there is a silvery dusted spot on the mesopleura, not on the sternopleura as
stated by Engel for costalis. S. costalis is the species recorded by Engel from
New Guinea.

Locality: Kuranda, Qld. (F. P. Dodd).

I accept the identification on the specimen as correct.

Tribe Rutiliini.

Brauer and von Bergenstamm used the name Rutiliidae for a group which
included only the genus Rutilia Robineau-Desvoidy. In a recent paper Dr. Engel
has dealt with “Rutiliidae sensu lat.”’, including in his treatment 11 genera, and
amongst them Amenia and Stilbomyia.

I consider the group dealt with by Engel to be heterogeneous, but several of
the genera he included are unknown to me at this time so that it is not possible
for me to allocate them definitely.

Townsend in 1916 designated vivipara (Fabricius) Robineau-Desvoidy as the
genotype of Rutilia. Engel cited desvoidyi Guerin as the genotype in his paper
and gave as the same species vivipara Robineau-Desvoidy. I consider these species
distinct, and as desvoidyi was not originally included it can not be accepted as the
genotype even if it were the same as vivipara, which I cannot admit is possible.
In the following pages I discuss the identity of these species.

In the collection of the United States National Museum there are several
species which are referable to Rutilia, and from an examination of these I have
drawn the following as evident generic characters: Suprasquamal ridge with
numerous long hairs, often curled and very conspicuous; centre of declivous area
above these hairs bare; face with a broad central vertical carina; arista pubescent;
abdominal sternites covered by the lateral margins of the tergites, sometimes
narrowly visible at apices; third and fourth visible tergites each with apical
central bristles, except sometimes the fourth in the male, and rarely the second
with a few shorter stout central apical bristles. In most of the Rutiliini before
me there is a very evident vein-like linear weak mark between the fifth and sixth
wing veins which simulates a vein, but this is not confined to this group and
recurs in Dexiinae of various groups.

My deductions as to specific characters hereinafter given are predicated upon
an examination of the species in the United States National Museum, no material
having been received by me from Australia direct. As is only to be expected in

346 NOTES ON AUSTRALIAN DIPTERA, Xii,

the case of such large, and usually brilliantly coloured, species there have been
many descriptions placed on record, and unfortunately it is very difficult to decide
definitely just how many valid species there are, and also to decide what species
are intended by certain of the descriptions. An extensive collection of the group
will undoubtedly provide a basis for a decision as to the number of species
involved, but only an examination of the type specimens will definitely decide
the exact specific names applicable to each, and when the type specimens are
scattered as in this case the work is too great to be undertaken by any one
individual.

RUTILIA VIVIPARA (Fabricius).

In the United States National Museum collection there is a specimen labelled
by Coquillett as this species. I have carefully gone over the specimen with the
original description and believe that there is a great probability that it is correctly
named. Wiedemann did not see the type specimen of this Fabrician species and
I do not know if it is in existence. There are, however, two other specimens
mentioned below which appear to confuse the issue somewhat and it will require
eareful work by someone on the ground to discover to what extent, if any, the
species varies in the characters mentioned below.

The specimen bearing Coquillett’s labei has the following characters: Hind
margin of the mesonotum with 4-6 strong bristles; prescutellar and presutural
acrostichal bristles well developed, 4 + 3-4 pairs; scutellum slightly convex, with
about 14 marginal bristles and a number of short but evident discals; supra-
squamal ridge with a few distinct hairs; second abdominal tergite with six short
stout central apical bristles; parafacials haired only on upper part to about apex
of second antennal segment; both spiracular coverings testaceous; submedian
thoracic vittae divergent behind, not continued much beyond suture; prosternum
and centre of propleura haired; anterodorsal bristles on hind tibiae irregular in
female.

Locality: Leura, N.S.W. (W. W. Froggatt). One female.

One specimen from Mittagong has two bristles on middle of apical margin of
second abdominal tergite, and has no presutural acrostichals, and two or three
pairs of prescutellars. The suprasquamal ridge is more pronouncedly haired than
in the above one.

Another specimen has no bristles on second abdominal tergite, but is very
similar to the preceding specimen in other respects.

It appears to be pertinent to note here that the presence of bristles on the
central apical portion of second abdominal tergite has been accepted as important
in distinguishing genera in this group, and if it is a variable character, which is
not at all unlikely, from an examination of the above three specimens, its value
is much less than indicated by Engel. Of course there may be more than one
species in the material before me, but it is impossible to be certain of this

without access to a larger series of specimens, and in better condition than those
now available.

RUTILIA DESVOIDYI Guerin.

Specimens accepted as this species by 'Townsend possess the following
characters: Parafacials haired to lowest margin of eyes; mesonotum with 4-6 hind
marginal bristles; only prescutellar acrostichal bristles present; no pronounced
discal bristles on scutellum, and 8-10 marginals, the disc subconvex; no distinct

BY J. R. MALLOCH. F 347

bristles on apical margin centrally of second tergite; male with the series of

bristles on antero-dorsal surface of hind tibia quite regular and close; in other

respects as the two last mentioned specimens of the preceding species.
Mittagong, N.S.W. (W. W. Froggatt); two males with label Australia only.
This species is decidedly more metallic green tinged on dorsum than is the

one accepted as vivipara.

RuTILIA INORNATA Guerin.

This species is usually much darker in colour than vivipara, being black, or
fuscous, with black legs, fuscous calyptrae, a dark mark near bases of wings,
and the dorsum of thorax and abdomen more or less bluish or violet coloured,
especially noticeable on the scutellum and abdomen.

The hind margin of mesonotum has a transverse series of 8-10 strong bristles,
the outer one on each side being the posterior dorsocentral; the scutellum is
flattened above, slightly angulate at apex in middle, and has about 14 strong
evenly spaced bristles, and no outstanding discals. The second abdominal tergite
has no central apical bristles, and the suprasquamal ridge is densely long haired;
while the parafacials are haired only to about apex of second antennal segment.

Locality: Queensland.

RUTILIA SPLENDIDA Donovan.

A brilliant metallic green or blue-green species. Head orange-yellow, upper
occiput green, the postocular orbits white dusted above; hairs on cheeks yellow.
Thoracic dorsum with four narrow black vittae, conspicuously white dusted
between the anterior portions of the vittae, on humeri, two lateral presutural
spots, and on an elongate spot above wing base; scutellum blue-green; mesopleura
with a large white dusted spot; pleural hairs black. Abdomen with a broad
black-blue dorsocentral vitta, and an apical fascia on second and third visible
tergites of same colour which widens out about midway to lateral curve, and
usually connects with a narrower basal fascia at that point leaving a large
submedian spot of green on each side of each tergite. Legs black. Bases of
wings blackened to apices of basal cells except for the usual central hyaline
spot. Calyptrae whitish at bases, brown beyond.

- Frontal lunule with some hairs; parafacials haired to below middle of eyes;
mesonotum with four or six posterior marginal bristles, the presutural acrostichals
small, but usually quite evident; postalar callosity with 4-5 bristles; sterno-
pleurals 2-3 + 1; scutellum slightly convex, with about 12 marginal bristles, and
some short bristles along edge of disc. Bristles at apices of third and fourth
tergites not very long. Anterodorsal fringe on hind tibia quite regular in both
sexes.

Locality: Cairns, N. Qld.

RUTILIA FORMOSA Robineau-Desvoidy.

A brilliant metallic coloured species. The thorax is usually pale blue-green,
with a slightly opalescent shade because of the presence of pale dusting, and the
abdomen is more emerald green, with a coppery tinge. The dorsum of thorax
has two faintly indicated dark submedian vittae anteriorly which are divergent
behind, and there are no distinct white spots present. The abdomen has a blue-
black dorsocentral vitta, and on apices of second and third visible tergites a
rather broad fascia of same colour which becomes obsolete at the lateral curve,

and is not present below. Head orange-yellow, frontal orbits and parafacials paler,
R

348 NOTES ON AUSTRALIAN DIPTERA, Xii,

becoming whitish along eyes, the orbits green at upper extremities; occiput and
the raised part of cheeks pale green, white dusted, postocular orbits white dusted;
occipital and genal hairs yellow. Legs black. Wings black near bases. The fine
thoracic hairs yellow. ,

Thoracic bristling as in splendida. MParafacials haired to, or almost to,
lowest level of eyes.

Localities: Townsville and Cairns, North Qld.

A female specimen from Shoalhaven has the third antennal segment black
except at base, but is otherwise the same as the above specimens.

A specimen from the old Fitch collection in the United States National
Museum, named by Bigot, agrees with the description given above.

In addition to the above-mentioned specimens I have one from New Guinea,
sent to me by Dr. Karny, which has the thoracic and cephalic hairs black, and
the basal dark mark on the wings very faint; and another female, from China,
belonging to the U.S. National Museum, which agrees very well with the one
from New Guinea, except that the hairs on occiput are yellow, except those on
margins above.

These variations may represent distinct species, but I have no males. —

I have carefully examined the two specimens in the United States National
Museum that were accepted as formosa by Townsend, and used as a basis for his
genus Chrysorutilia. These are larger and more robust than the specimens dealt
with above, and are darker in colour, with the genae much less conspicuously
green, and the face below carina, and the third antennal segment entirely fuscous.
In addition to these characters the female has a quite conspicuous apical central
depression on the fourth visible tergite as in typical species of Rutilia, which is
not the case in the other females before me.

In my opinion Chrysorutilia is not a good genus, and the genotype is not
correctly identified if Engel’s identification is correct, the latter being the form
with the yellow thoracic hairs first listed in this paper.

I have examined several additional species of Rutilia in the United States
National Museum collection, but at the time of writing cannot be absolutely certain
of the specific identity of any of them except the following two, so leave them
aside temporarily.

RUTILIA ERICHSONI Engel.

Head yellowish fulvous, occiput and raised part of cheek brassy green,
occiput, genae, and parafacials, yellow haired. Thorax violet-black, with black
hairs and the following bluish or greenish silvery marks: Two presutural sub-
median dorsal vittae, four spots on each lateral margin of mesonotum, the middie
pair one above the other, and a spot on mesopleura. Abdomen coloured as thorax,
second and third visible tergites each with an anterior marginal, metallic green,
or blue-green, centrally interrupted, fascia, fourth tergite with a black central
spot on anterior margin, the remainder of surface metallic coppery green. Wings
dark at bases. Calyptrae yellowish white.

Parafacials haired to lowest level of eyes. Scutellum convex, with eight
marginal, and some short discal, bristles. Second tergite of abdomen with a pair
of short apical central bristles, fourth tergite weakly bristled, not concave at apex.

Length, 13 mm,

Locality: Waroona, W.A., 25 Jan., 1909 (G. F. Berthoud).

BY J. R. MALLOCH. 349

RUTILIA ARGENTIFERA Bigot.

Head as in erichsoni. Thorax violet-black, posteriorly and on scutellum
markedly more bluish, the presutural pair of vittae vestigial and narrow, each
lateral margin of mesonotum in the specimen before me with but two bluish
white marks, and the mesopleura with one. Abdomen violet-black, second and
third tergites each with four round greenish or bluish silvery spots, fourth with
a large silvery white spot on each side. Wings blackened at bases. Lower calypter
browned apically. Thoracic hairs black.

Structurally similar to erichsoni.

Length, 13 mm.

Locality: Cairns, North Qld.

There are a few slight discrepancies between the above descriptions and that
of this species given by Engel, but they are not important enough to justify a
belief that they do not belong to the same species.

I present below a key to the foregoing species.

Key to the Species.
1. Parafacials not haired below middle of eyes; species not bright metallic green nor

ITS SSE OT oa Por. eee San Manic steuey eecaerretay suchen telist Median sitar enaie tara arcenenetts Yorenel Guntettsesvemer eneweleoteits 2
Parafacials haired to below middle, often to lowest level of eyes; if only to middle
themspeciesmas!) larsely emetallicwereentec-eierroeiacicecieietelcy ct oiclonetelcliclcnesoietetoheteleicier 3

2. Hind margin of mesonotum with 4-6 strong bristles; scutellum subconvex ........
BIO COTO EDICT CIES De OREO aor miad © eas cic Or OIcL CIO TerG Rony OnERCD O mnCR Or oto IG aa vivipara (Fabricius)

Hind margin of mesonotum with 8-10 strong bristles; scutellum flattened on disc
STS SUMS ER ONO Oe OU REE RRR Cl RCHSEE OURO MER CRC CoE CCR SHER CHC RRR ED eee Eh nr RSS inornata Guerin.

3. Thorax fuscous, with slight greenish or bluish metallic tinge, and no conspicuous
white markings on either it or abdomen ...................... desvoidyi Guerin.
Thorax either brilliant metallic green or blue-green, if black then with conspicuous
white markings, and with white or green markings on abdomen .......... 4

4. Thorax brilliant metallic emerald green, without white markings .. formosa Rob.-Desv.
Thorax either metallic blue-green, or black, with metallic reflections, and always
WH COMIMCHOUE Whee, ie MOINES oo 5c0casoonocu5 db odDoDD Un UOUGaDOOHOoNONE 5

5. Thorax and abdomen brilliant metallic blue-green, the former with four narrow
blackish vittae anteriorly and between these the ground colour is quite
conspicuously obscured by white dusting, lateral margins of mesonotum each with

four white spots, the median pair one above the other; abdomen with black

basal and apical fasciae on second and third tergites which are usually connected
midway from the central black vitta to lateral curve so as to leave two large

green spots on each side of each of the tergites .............. splendida Donovan.
LhHonaxwandsapdomentviolet-blackerermeirerecireiicrerlers cucncicrcicloreislsicisicheleiecckcusteieien cachet ons 6

6. Abdominal tergites 2 and 3 each with four rather small round bluish, or greenish,
silvery spots on disc, the fourth with two larger white spots .. argentifera Bigot.

Sides of first tergite, and anterior margins of second and third on sides, broadly
metallic green, fourth tergite metallic green, with a central black spot on
AMLECTION PMLA ETN so orca. oy lish sh este cnesee eR Me suclione_cenorsic ston enaisiaerey enero) cereus erichsoni Engel.

N.B.—This key should be used carefully in connection with the foregoing notes

on the included species to prevent errors in identification.

Genus AMPHIBOLIA Macquart.

This genus is similar to Rutilia in having the suprasquamal ridge haired, and
the declivity above these hairs bare. The specimen of the genotype, valentina
Macquart, which I have examined, has the second and third visible tergites of
abdomen each with two small bristles near anterior margin in centre, and some
others near apex in centre. The parafacials are bare below middle, and there are
but three strong bristles on the postalar callosity.

I have seen only one species referable here, the other one included by Engel,
speciosa Erichson, being a species of Formosia.

350 NOTES ON AUSTRALIAN DIPTERA, Xii,

Genus FormostA Guerin.

This genus as included in Engel’s paper may be heterogeneous, but my
definition of it is based upon mirabilis Guerin and flavipennis Macquart, the
former being the genotype. It is distinguished from Rutilia by the lack of hairs
on the suprasquamal ridge, and the presence of long hairs on centre of the
postalar declivity. The arista is also often much more distinctly haired, the
posterior margin of mesonotum and of second abdominal tergite in centre are
more strongly bristled as a rule; and the prosternum is bare, while the centre of
the propleura is haired. In addition to these characters the species before me all
have a series of very prominent erect spines on the apical margin of the tergites
below, and the parafacials bare below bases of antennae. The colour of the
parafacials cited by Engel as a distinguishing character of the genus is valueless,
in my opinion, though it appears to distinguish the two groups here included,
those with the distinctly haired aristae having a green colour while those with
the pubescent aristae have no green colour present on the parafacials.

FORMOSIA FLAVIPENNIS Macquart.

This species differs from its congeners of metallic green colour in having the
legs fulvous yellow, only the apices of the tarsi being infuscated, and the pleural
hairs entirely yellow. The thorax is emerald green, with the sutures fulvous
yellow, and dorsum faintly quadrivittate anteriorly; the head is fulvous yellow,
with occiput, frontal orbits, and the greater part of parafacials and cheeks,
opalescent green. Abdomen fulvous, first tergite above, and apices of other
tergites black, disc of second tergite and sides of third and fourth, coppery red,
disc of third and fourth tergites largely blue, sides of second green. Wings
yellow at bases, not darkened; calyptrae and halteres yellow.

Arista with the longest hairs about as long as width of third antennal segment.
Only the posterior dorsocentral bristles present; sternopleural 1; scutellum
flattened above, broadly subtriangular, with about 14 irregular marginal bristles.

Locality: Cape York, N. Qld.

FORMOSIA MIRABILIS Guerin.

This species, as identified by Townsend in the United States National Museum,
is very similar to flavipennis, but the legs are fuscous, and the hairs on mesopleura
are black. The thoracic dorsum has four quite distinct black vittae on the
metallic emerald-green ground colour. ‘The abdomen is coppery green, black on
first tergite, a median vitta on all tergites which is wider behind, the apices of all
tergites, with a large spot-like enlargement on each side of each tergite, the black
apical margin continued below, and becoming rather abruptly linear near lateral
margin. Wings hardly darkened at bases. Calyptrae brownish yellow.

Eyes of male separated by not over the width of third antennal segment; arista
rather long haired.

Locality: Aru Island.

This specimen looks like a dark example of flavipennis, but may be distinct.
The apex of abdomen is paler than in the above female, instead of darker which one
would expect to be the case if they are the same species.

FORMOSIA ATRIBASIS (Walker).
This species was included in Rutilia by Engel, but apart from the pubescent
arista there is no character of any import that one can use for its separation
from Formosia as herein accepted.

BY J. R. MALLOCH. 351

It is readily separated from the preceding two by the fuscous head, which has
dense whitish dust everywhere except on the interfrontalia, instead of being green,
the black thorax, which is greenish tinged and quadrivittate on dorsum, the
presutural portion being quite distinctly white dusted, and has a large white
dusted mark on the mesopleura. The abdomen is black, with second tergite green,
conspicuously reddish coppery tinged, and with a large transverse black mark at
bases of the apical central bristles, third tergite coppery green on anterior margin
each side of the black central mark. Legs, bases of wings to apices of basal cells,
and the calyptrae, deep black. )

Arista pubescent. Dorsocentrals 3 + 5, the anterior two pairs of postsuturals
weak, presutural acrostichals weak but evident; scutellum subconvex; sterno-
pleurals 1 + 1, anterior one weak.

Locality: Kuranda, Qld. (F. P. Dodd).

ForRMOSIA SPECIOSA (EHrichson).

This species agrees with atribasis in having the arista merely pubescent, but
differs markedly in colour, entirely lacking metallic tinge, being entirely black,
with markings of dense greyish white dust on thorax and abdomen. Head
testaceous, frons and upper occiput fuscous, all except the interfrontalia densely
grey dusted; third antennal segment blackish except at base; palpi testaceous
yellow. Thorax opaque black, with the following white dusted marks: four vittae
and a small anterior central mark in front of suture, four spots behind suture,
a spot on mesopleura and another on sternopleura. Abdomen densely whitish grey
dusted, black on the following parts: first tergite, a transverse spot near apex
of second tergite in centre, a cordate central spot and a much larger one on each
lateral curve of third tergite, all of which connect with larger spots on fourth,
and a spot on extreme lateral margin of each tergite below. Legs fulvous yellow.
Wings slightly infuscated at bases. Pleural hairs yellow.

Thoracic bristling as in atribasis, but the presutural acrostichals lacking.

Locality: Wickham, N.S.W.

This is the genotype of Huamphibola Townsend, but I do not consider it
entitled to generic separation from Formosia, the only character distinguishing it
being the pubescent arista. It may ultimately be accorded subgeneric rank with
atribasis. I have seen several other species that would also fall in the group, but
cannot now identify them.

Genus PRODIAPHANIA Townsend.

This genus is distinguished from all three of this tribe previously dealt with
in this paper by the absence of hairs on the suprasquamal ridge and postalar
declivity. Other distinguishing characters lie in the extremely small palpi, all
of the others having these organs long and slender, and the long narrow upper
calypter, which is as long as the lower one and has some small black hairs
invading the disc basally on the outer half. The arista is long haired, and the
postalar bristles normally four in number. The conspicuously explanate costa is
not confined to this genus, as I have before me a species of Rutilia which very
closely resembles in superficial appearance the only known species of Prodiaphania,
and has the same costal character.

Prodiaphania has the prosternum bare and the propleura haired in centre,
and except for the peculiar upper calypter is closely similar to some genera of
Dexiinae. I had thought it might be possible to use this lengthening of the upper

352 NOTES ON AUSTRALIAN DIPTERA, Xii,

calypter as a group character, as it is evident in Rutilia and Formosia, but it is .
not so marked in either of these genera, and is slightly variable in the different
species.

The only species of Prodiaphania is testacea Macquart, which I have seen from
Cairns, North Queensland.

Of the other genera dealt with by Engel I have not seen the following:
Paramenia Brauer and von Bergenstamm; WMicrotropeza Macquart, Chrysopasta
Brauer and von Bergenstamm, Senostoma Macquart, and Paramphibola Brauer
and von Bergenstamm.

Judging from the descriptions the first two do not fall within the group as
accepted in the present paper, having no facial carina, and without an examination
of the others it is impossible for me to indicate their relationships.

In the United States National Museum there are several genera placed in this
group by Townsend which were not mentioned by Engel, and some of these are
properly referable here, though one or two are not. I have already referred to
Chrysorutilia and Euamphibolia, and below deal with some of the other genera
which may be considered as referable to the group or in its immediate vicinity.
I do not guarantee the correctness of the specific identifications given, as that
must await type examination.

Genus GRAPHOLOSTYLUM Macquart.

I have not seen dorsomaculatum Macquart the type species of this genus, but
in the United States National Museum there is a specimen identified by Townsend
as decorum Guerin which ‘is placed in this genus. This specimen agrees with one
other so named by Bigot in the same collection. Engel places decora as a synonym
of splendida Donovan, but the two specimens before me are quite different in the
colour and markings of the abdomen from the series identified as splendida. The
general colour of the abdomen is a deep bluish black, on the disc of tergites 2 and
3 there is a pair of small round green spots, on the lateral curves of tergites 1 to 3
there is a larger transverse green mark, and on each side of fourth tergite there
is a large green mark, the margins of all of these markings suffused with Prussian
blue colour.

Parafacials haired to lowest level of eyes.
Locality: Sydney, N.S.W.
This species is a genuine Rutilia, but dorsomaculatum may not be.

In the key to the species of Rutilia given on a previous page in this paper the
present species will run down to splendida, but the smaller size of the dorsal spots
on abdomen, and the greater proportion of the parafacials which is haired will
readily distinguish the two forms, which may not be distinct species but only
varieties of one.

Genus RutTiLopEx1A Townsend.

This genus is doubtfully referable to this group. The face in profile is more
protuberant than in Rutilia; postalar declivity, suprasquamal ridge, and prosternum
bare; centre of propleura with hairs; parafacials bare below bases of antennae;
arista pubescent. Calyptrae as in typical Dexiinae. Postalar bristles 4-5. Abdomen
as in Rutilia, with no discal bristles except on fourth tergite. The genotype is
identified as angustipennis Walker.

BY J. R. MALLOCH. ; 353

Genus CHAETOGASTER Macquart.

This genus is similar to the preceding one, but has quite distinct presutural
acrostichal bristles on thorax, and the abdominal tergites 2-4 have each one or more
pairs of quite strong discal bristles. Facial carina not so prominent. Genotype,
violacea Macquart.

Occurs in Victoria and New South Wales.

Genus CHLOROTACHINA Townsend.

This genus has no facial carina and does not come within the scope of the
group as accepted herein. It would run to Microtropeza Macquart in Engel’s key
to the genera if one cared to consider it as belonging to “Rutiliidae sensu lat.’’, but
the third antennal segment is much longer than the second, and the insect is a
bright metallic blue-green in colour, two characters which exclude it from Engel’s
concept. The genotype is flaviceps Macquart.

Locality: Cairns, North Qld.

DESCRIPTIONS OF NEW SPECIES OF AUSTRALIAN COLEOPTERA.
PART Xix.
By Arruur M. Lea, F.E.S.
[Read 27th July, 1927.]

The species dealt with in the present paper all belong to the family
Curculionidae.

CorpTORHYNCHUS ALBIVARIUS, 0D. Sp.

Black, legs sometimes obscurely reddish. With numerous white spots and
patches of scales, and many white setae.

Head with numerous punctures in front, with a somewhat horseshoe-shaped
impression between eyes. Rostrum slightly dilated to between antennae; with
crowded naked punctures in front. Antennae long and thin. Prothorax about as
long as the greatest width, sides strongly rounded, with dense punctures, each
with a white seta. Elytra ovate, strongly convex, greatest width about twice that
of prothorax; with rather shallow striae containing large punctures. Legs long,
femora clavate. Length, 3-5-5 mm.

Habitat. Queensland: Coen River (W. D. Dodd and H. Hacker).

In appearance close to C. jansoni, but the vitta on each side of the base of
pronotum subtriangular in shape with the point in front; it is usually connected
at the base with a longer and wider one, which touches the coxa and is curved
upwards in front so that its tip is visible from above. The rostrum is clothed
with white scales, on some specimens appearing as two lines, each of which is
dilated near an eye. On the elytra the spots vary considerably, often on the
sides of an individual; there is usually a semi-double one on the suture just
beyond the middle (absent from one specimen), on the third interstice there are
one, two, three, or four round ones, and a long one near apex (on the right
elytron of one specimen there are seven spots on the third interstice), on the
fifth there are one or two round ones about the middle, a long one at base and
another beyond the middle; the seventh and ninth usually have two spots on
each, and the lateral interstice is usually narrowly white. There is a white
spot on each side of the mesosternum and metasternum, and other white scales
on the under parts.

CoproRHYNCHUS OCULARIS, Nl. SDP.

Black, antennae and usually parts of the legs obscurely reddish. Clothed
with white setae and numerous spots of white scales.

Head depressed between eyes, these unusually prominent. Rostrum suddenly
elevated above head at base, in front with naked punctures and a short median
line. Antennae long and thin. Prothorax distinctly longer than wide, sides
strongly rounded; with dense punctures each containing a seta. LEHlytra ovate,

BY A. M. LEA. 355

strongly convex, greatest width about twice that of prothorax; with rows of large
punctures, mostly wider than interstices, becoming smaller and in deeper striae
posteriorly. Legs long, femora clavate. Length, 4-5 mm.

Habitat.—Queensland: Darnley Island (H. Elgner and A. M. Lea).

The base of the rostrum appears elevated and subtriangularly produced
between the eyes; the latter from behind appear to be separated from the head
by slight notches; the prothorax is longer than is usual. The white scales on the
best preserved specimen are fairly dense on the basal two-thirds of rostrum and
between the eyes, form a short basal vitta on each side of the pronotum and are
dense on the sides (but invisible from above); on each elytron (as seen from the
side) there is a short basal fascia (close to two large spots on the under surface),
another before the middle (composed of three conjoined spots, one on each
interstice), and a subtriangular patch beyond the middle; at the apex there is
a short vitta on the third interstice, connected with a narrow marginal stripe,
there are several feeble spots elsewhere, including one on the suture. On three
other specimens the markings are less defined, partly due to abrasion. On another

the white scales are much more numerous, but the markings are less sharply
indicated.

CoPTORHYNCHUS TRIVITTATUS. Nl. SD.

Black, parts of antennae and of legs obscurely reddish. Clothed with white
setae and spots of white scales.

Head with numerous punctures in front, with a small interocular fovea.
Eyes prominent. Rostrum in front with numerous naked punctures, and a median
depression. Antennae long and thin. Prothorax moderately transverse, sides
strongly rounded, base very little wider than apex; with crowded punctures,
each containing a seta. Elytra ovate, strongly convex, almost twice the width
of prothorax; with rows of large, deep punctures, much wider than interstices,
even posteriorly. Legs long, femora clavate. Length, 4-5-5 mm.

Habitat.—Queensland: Darnley Island (H. Elgner).

The base of the rostrum and the eyes are very different from those of the
preceding species. The white scales are dense on the basal two-thirds of rostrum
and around the eyes; on the pronotum they form three basal vittae and a stripe
adjacent to each coxa, with its ends turned upwards. On the elytra they form
two spots on the suture, four on the third interstice (and a large spot near
apex where several interstices join), three on the fifth, three on the seventh
and three or four on the ninth, with the margin narrowly white from near the
base. The sides of the under surface from the mesosternum to the apex of the
first abdominal segment are densely clothed with white scales; the middle of
the first segment is slightly depressed, with long thin setae. The type is probably
a male, a second specimen has the abdomen evenly convex, with the sides of the
first segment less densely clothed; some of the spots on its upper surface are less
defined than on the type.

RHINOSCAPHA INTERRUPTA, N. Sp.

¢. Black. Sparsely clothed with inconspicuous scales, except on elytra, on
which they form distinctive markings.

Head with sparse and minute punctures. Rostrum almost the length of
prothorax; median groove deep and single at base, becoming trifid in front;
sublateral sulcus rather deep, both ends closed. Antennae rather thin. Prothcrax

s

356 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

slightly longer than wide, widest at apical third, somewhat depressed along middle ~
but with an irregular carina, sides vermiculate; with small scattered punctures.
Elytra elongate-subcordate, base bilobed, sides widest about middle; with regular
rows of large punctures, the interstices in places faintly wrinkled and with sparse
minute punctures. Legs rather long. Length (excluding rostrum), 14-19 mm.

©. Differs in having the rostrum slightly stouter, prothorax slightly wider,
elytra much wider, abdomen more convex and legs somewhat shorter.

Habitat——Queensland: Murray Island (A. M. Lea), Darnley Island (C. T.
McNamara).

On each elytron there are several spots or patches of white scales having a
distinct greenish gloss, and usually mixed with a reddish meal: two near base,
of which the inner one is slightly oblique, and about half the length of the
prothorax, and a smaller one behind the shoulder, and partly visible from above;
on the fourth interstice before the middle there is a short vitta, behind this an
oblique one on the third, and level with the apex of this a smaller one on the
second, towards the side there is a small one on the seventh and eighth, on the
fifth there is a distinct one on the apical fourth; the marginal interstice from
near the base, and the next two near apex are densely clothed. On each side of
the prothorax, but invisible from above, there is a vitta of whitish scales with a
greenish gloss. The specimen from Darnley Island is a large female, with the
elytral markings white and slightly more extended. Its nearest ally before me
appears to be R. tricolor, from New Guinea, on which the subbasal and submedian
markings on the elytra are connected, and which has distinctive markings on
the sides of the prothorax.

MANDALOTUS.

In my recent revision of the genus I stated! of the apparently allied genus
Timareta, “Several species with ocular lobes’. This should have been “without’’.

MANDALOTUS PYRIFER Lea.
Rec. S. Aust. Museum, 1926, p. 167.

A female, from the Victorian Alps, in the National Museum, Melbourne,
evidently belongs to this species. It differs from the type male in having the
mesosternum without the intercoxal process produced, two basal segments of
abdomen slightly convex, and not setose, front tibiae without distinct teeth, and
hind ones less curved.

MANDALOTUS MIRABILIS Lea.
Trans. Roy. Soc. 8. Aust., 1907, xxxi, p. 152.

Four males, in the National Museum, Melbourne, from the Howitt collection,
belong to this species, but the long hairs on the under surface are plastered down
on three of them, so that the character given in the 1926 key of the genus:
“Under surface with dense and long hairs” does not appear to apply to them. Hach
middle coxa is seen to have an obtuse tooth (the end of the arcuate carina).
M. campylocnemis, somewhat similarly armed, has the middle coxae more than
twice as wide apart, and very different legs; M. valgus is a smaller species, with
shorter and sparser clothing on the under surface.

1Lea, Rec. 8S. Aust. Mus., 1926, p. 148.

BY A. M. LEA. 357

Two females, associated with the males, apparently belong to the species;
they differ in being smaller, sides of prothorax more evenly rounded, elytra more
parallel-sided, metasternum and abdomen flat along middle, middle coxae unarmed
and tibiae simple. On one of them there are four conspicuous pale spots on the
head: a small one margining each side, a still smaller one at the base of the
rostral carina, and a large medio-basal one; on the other female and on the males
the spots are feeble or scarcely traceable.

MANDALOTUS STERNOCERUS, DN. Sp.

6. Black, antennae tarsi and parts of tibiae reddish. Densely clothed with
muddy-brown scales, interspersed with setae.

Rostrum with median carina concealed or absent. Antennae fairly long and
thin. Prothorax moderately transverse, sides evenly rounded, with numerous
small rounded granules traceable through clothing. Elytra wider than prothorax,
base almost evenly arcuate, with rows of large punctures, much wider than
interstices, but appearing much smaller through clothing, alternate interstices
slightly elevated and in places subtuberculate. Mesosternum with a five-sided
projecting intercoxal process, widest at about one-third from base, the apex
truncated and about half the width of base. Metasternum and abdomen each
with a basal row of large punctures, and with a wide shallow depression continued
on to second segment. Front coxae widely separated, front tibiae with a large
obtuse tubercle near base, the sides flanged and apex armed; hind tibiae with a
fairly large tooth near base, between it and apex shining and multicarinate.
Length, 5-6 mm.

. Differs in being slightly wider, mesosternum unarmed, two basal segments
of abdomen evenly convex, tibiae unarmed near base, and the hind ones not
multicarinate.

Habitat.—Victoria: Types in National Museum, Melbourne.

Another species of the genus with remarkable tibiae. In the 1926 key
of the genus’ it could be associated with M. intercoxalis, from which it differs
in the smaller size of the intercoxal process and the remarkable tibiae;
M. carinatipes has the hind tibiae multicarinate, but the front tibiae are very
different and the mesosternal process is shorter and broader. The front tibiae
from some directions appear to have an acute apical tooth, projecting downwards
about the same length as the subbasal one, but from other directions it is seen
to be a thin flange, with its apex semidouble. On the elytra the setae are more
numerous on the odd than on the even interstices, and a few are clustered together
in places, but hardly enough to form fascicles, at the base on the male they form
two rather pale spots; there is an oblique series of feeble elevations crowning the
apical slope, but the only distinct tubercle on each elytron is one on the side near
the shoulder.

od

MANDALOTUS ARMICOXIS, N. Sp.

6. Black, antennae and tarsi obscurely reddish. Densely clothed with sooty-
brown scales sparsely interspersed with setae, but the latter becoming longer
and more numerous on legs:

Rostrum with median carina concealed or absent. Antennae moderately long
and thin. Prothorax moderately transverse, roughly and irregularly granulate.
Elytra slightly narrower than widest part of prothorax, base trisinuate, with rows

1Lea, Rec. S. Aust. Mus., 1926, p. 149.

358 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,
of large partially concealed punctures, third, fifth and seventh interstices —
irregularly elevated or subtuberculate, a distinct tubercle on the side behind each
shoulder. Metasternum and abdomen shallowly concave and each with a basal
row of large punctures. Front coxae widely separated, very large, semidouble,
the hind part of each with a conspicuous subconical tubercle; front tibiae strongly
curved at apex; hind tibiae less strongly curved but still conspicuously so.
Length, 5-5-5 mm.

Habitat.——Victoria: Two males in National Museum, Melbourne.

In the 1926 key of the genus, this species could be associated with M. valgus,
but it is readily distinguishe@ from that species, as from all others, by the front
coxae, these being of unusual size, semidouble and conspicuously tuberculate.
The front trochanters are also peculiar; from one direction they appear to be
transverse and to isolate completely the coxae from the femora, although from
another direction the femora are seen to junction with the coxae.

BaARIS BITUBERCULATA, Nl. SD.

Black, shining. With sparse white setae, nowhere forming distinct spots.

Head with fairly dense but not very sharply defined punctures; with a small
interocular impression. Rostrum evenly curved; punctures more sharply defined
than on head, although no larger. Scape inserted about two-fifths from apex of
rostrum in female, slightly nearer in male. Prothorax almost parallel-sided except
near apex, which is suddenly narrowed; with dense punctures of moderate size
on sides, becoming smaller and somewhat sparser in middle, but quite as sharply
defined. Elytra elongate-subcordate; with narrow striae containing small
punctures, but becoming larger about base; interstices with very minute punctures,
but some fairly distinct ones about base and apex (where also the derm is
subopaque); a small preapical callus on each side just before the junction of the
third and sixth interstices. Sterna with crowded punctures about as large as
those on sides of prothorax, becoming smaller and sparser on abdomen. Femora
acutely dentate. Length, 3-5 mm.

Habitat— Queensland: Murray Island (A. M. Lea).

Belongs to Section 1! of the genus, but distinct from all other species of the
subfamily (except B. niveodispersa) by the elytral tubercles. The groove in front
of the prosternum can be seen only after the head has been removed. From the
sides there appears to be a shallow notch between the head and rostrum.

BaRIS COELESTIS, nl. Sp.

Deep metallic blue. Setae minute and inconspicuous.

Head with minute punctures, with a shallow interocular impression. Rostrum
strongly curved in a line with that of the head; with small punctures more
distinct than on head, sides opaque. Antennae inserted about one-third from
apex of rostrum. Prothorax moderately transverse, sides almost evenly decreasing
in width from base to apex; with rather small and not very dense, sharply defined
punctures, becoming very small in front, on the sides larger and more crowded,
but an impunctate polished space on each side. Elytra with sides very feebly
decreasing in width from shoulders to near apex; with narrow striae containing
small deep-set punctures, distinct only about basal fourth; punctures on interstices
scarcely visible. Metasternum with crowded, sharply defined punctures, becoming

1 Lea, Trans. Roy. Soc. 8S. Aust., 1906, p. 84.

BY A. M. LEA. 359

smaller and sparser on the rest of the under surface. Front femora very minutely
dentate, the others not at all. Length, 2 mm.

Habitat.—Queensland: Cairns district (F. P. Dodd).

A beautiful blue species. As the prosternum has a groove bounded by distinct
walls, it must be referred to Section 1 of Baris rather than to Ipsichora; the
groove, however, can be seen distinctly only after the head has been removed.
The legs are somewhat darker than the upper parts, and the under parts have a
slight greenish gloss; the antennae are almost black. Owing to the opacity of the
sides of the rostrum, its middle appears to be finely ridged; it is narrowed between
the eyes, these being larger than usual. The polished space on each side of the
prothorax is invisible from above. The male has a large, shallow, intercoxal
depression.

BARIS PULCHRIPARVA, DN. Sp.

Bright metallic blue, antennae blackish. Prothorax with a small spot of
white scales on each side of base, a similar spot on each side of metasternum,
elsewhere sparsely clothed or glabrous.

Head minutely punctate, with a small depression in front. Rostrum strongly
curved, with minute but rather sharply defined punctures. Antennae inserted
two-fifths from apex of rostrum. Prothorax gently narrowed from base to near
apex and then strongly to apex itself; with rather small but sharply defined
punctures, becoming larger and more crowded on sides. Elytra with sides
nowhere quite parallel, with narrow striae containing deep-set punctures, distinct
only on basal third; interstices very minutely punctate. Metasternum with
crowded punctures, about as large as on sides of prothorax, the abdomen with very
irregular ones, but mostly sparser and smaller than on metasternum. Femora
edentate. Length, 2:5 mm.

Habitat— Queensland: Cairns district (F. P. Dodd). Unique.

Another beautiful blue species of Section 1; readily distinguished from the
preceding species by the white spots on the prothorax and metasternum. The
rostrum from above appears parallel-sided, from the sides it appears to be
separated from the head by a feeble notch, to be angularly bent at the middle, with
the apex much narrower than the base. As its abdomen has a shallow depression
on the two basal segments the type is probably a male.

BARIS HOPLOCNEMIS, N. Sp.

Deep metallic blue, legs and antennae blackish. Upper surface glabrous, except
for a white spot on each side of base of pronotum; under surface sparsely clothed,
but a conspicuous white stripe on each side of metasternum.

Head with minute punctures, and with a shallow interocular depression.
Rostrum strongly and separately curved, with numerous small but sharply defined
punctures. Antennae inserted one-third from apex of rostrum. Prothorax almost
as long as wide, sides almost parallel to near apex; with numerous sharply defined
punctures of moderate size, becoming minute in front and larger and crowded
on sides. Elytra slightly wider than prothorax, parallel-sided to near apex; striae
narrow and apparently impunctate, interstices impunctate except close to base.
Under surface with crowded and coarse punctures, except on most of abdomen
and middle of metasternum. Legs rather long; femora edentate; front tibiae with
a conspicuous projection on middle of lower surface. Length, 2 mm.

Habitat Queensland: Cairns district (F. P. Dodd).

360 NEW SPECIES OF AUSTRALIAN COLEOPTERA, XiX,

A narrow, beautiful species, with distinctive front tibiae. The prosternal
groove is rather shallow, and its sides are not bounded by acute ridges in front,
but it seems desirable to refer it to Section 1, with the other blue species of the
genus. It differs from B. pulchriparva in being more parallel-sided, prothorax
less narrowed to apex, with larger punctures, metasternal episterna clothed
throughout, abdomen without a longitudinal impression and in the front tibiae.

A specimen, from Glen Lamington in the Stockholm Museum, differs in being
larger (2-5 mm.) and more greenish than blue. The abdomen of both specimens
is evenly convex, but the armature of the front tibiae is probably confined to
the male.

BARIS ORTHODOXA, Nl. SDP.

Black, somewhat shining, with a faint bronzy gloss; under parts with small
white setae, upper surface glabrous.

Head opaque and with very minute punctures. Rostrum strongly curved,
almost in a line with curvature of head; with small seriate punctures behind inser-
tion of antennae (at apical third), more sharply defined in front. Prothorax strongly
transverse, sides almost evenly narrowed from base to apex; middle with rather
small, sharply defined punctures, becoming larger and crowded towards and on
sides. Elytra cordate, outlines continuous with those of prothorax; with narrow
striae, containing small angular punctures, deeply set posteriorly; interstices each
with an inconspicuous row of punctures. Under surface with crowded punctures,
smaller and sparser on parts of abdomen than elsewhere. Femora edentate.
Length, 2 mm.

Habitat.—Queensland: Cape York. ‘Unique.

A small species of Section 1, slightly wider than B. vagans, and with the
pectoral canal more sharply defined, it is narrow and rather deep, but without
acutely ridged margins. As the basal segment of its abdomen has a distinct median
impression, the type appears to be a male.

BARIS PARVONIGRA, N. Sp.

Black, slightly shining, some parts opaque. Upper surface glabrous, under
parts with minute white setae.

Head minutely punctate. Rostrum strongly curved, in a line with curvature of
head; with fine ridges, alternated with rows of small punctures to apical third
(where the antennae are inserted), then with punctures only. Prothorax
moderately transverse, apex about half the width of base; with rather small
punctures in middle, becoming crowded and larger on sides. Elytra cordate, out-
lines not quite continuous with those of prothorax; with narrow striae containing
small, deep-set punctures, vanishing posteriorly; interstices each with an
inconspicuous row of punctures. Under surface with crowded punctures, some-
what sparser on abdomen than elsewhere; basal segment with a faint depression.
Femora edentate. Length, 1-75 mm.

Habitat—wNorthern Queensland (Blackburn’s collection). Unique.

A minute comparatively narrow species of Section 1, in general appearance
like B. subopaca, and with the prosternum similarly grooved. It is about the
size of B. vagans, but the prothoracic punctures are more crowded.

BARIS SETISTRIATA, N. Sp.
Black, tarsi obscurely diluted with red. Moderately clothed with white setae,
on the elytra forming a distinct row on each interstice.

BY A. M. LEA. 361

Head with minute punctures. Rostrum about the length of prothorax,
moderately curved, with rows of squamiferous punctures to middle (where the
antennae are inserted), in front of which the punctures are smaller, more
numerous and naked. Prothorax slightly transverse, sides almost parallel to
near apex; with dense and rather small punctures, becoming larger on sides. Elytra
suboblong, base bilobed, with narrow striae containing deep-set punctures, fairly
distinct about base only. Femora edentate. Length, 1-5 mm.

Habitat.——Queensland: Rockhampton (A. M. Lea).

A small, suboblong species, with a line of white setae on each elytrai
interstice; it belongs to Section 1, but the pectoral canal, although distinct in
front, is bounded by rather feeble ridges. On the male the antennae are inserted
slightly nearer the apex of the rostrum than on the female, and the base of
its abdomen has a slight longitudinal impression. The outlines are much as on
B. oblonga, but the elytral clothing does not conceal the derm on which it rests,
whereas on that species, there appear to be alternating white and black lines.
B. vagans is slightly larger, less parallel-sided, and has elytra without seriate
clothing. B. australiae is smaller and has more conspicuous clothing.

BARIS TRISINUATA, DN. Sp.

Black, parts of under surface and of legs obscurely diluted with red. Upper
surface glabrous, under parts almost so.

Head minutely punctate. Rostrum thick, strongly and equally curved; with
coarse, crowded punctures on sides, smaller and more sharply defined on upper
surface. Antennae inserted nearer apex than base of rostrum. Prothorax with
base almost twice as wide as median length, apex less than half the width of base;
with numerous rather small punctures, becoming larger and closer together on
sides. Elytra very wide, outlines subcontinuous with those of prothorax, sides
nowhere parallel, base distinctly trisinuate; with narrow striae, containing deep-
set punctures, but becoming round on basal half of two sutural ones; interstices
polished and impunctate, but about outer apex opaque and with small punctures.
Under surface with very variable punctures; crowded in front of prosternum, and
on apex of abdomen, almost absent from third and fourth segments. Femora
grooved, edentate, and coarsely punctate. Length, 3 mm.

Habitat.—North Queensland (Blackburn’s collection). Unique.

A short species of Section 1, wider in proportion than any species of the genus
previously recorded from Australia. The pectoral canal is wide and shallow, in
front it is somewhat deeper in the middle with feebly carinated sides. As the two
basal segments of its abdomen are depressed in the middle the type is probably
a male.

BARIS MELANOSTETHA, 0. SD.

¢. Castaneous-brown, scutellum, suture and metasternum black or blackish.
Upper surface with sparse white clothing but forming a small medio-basal spot
on pronotum; on under parts rather dense.

Head with small, crowded punctures, and a faint interocular depression.
Rostrum scarcely the length of prothorax, with fine ridges and irregular punctures
to apical third (where the antennae are inserted), in front with punctures only.
Prothorax longer than apical width, which is about half that of base; with rather
dense punctures of moderate size. Elytra with outlines subcontinuous with those
of prothorax, sides in part subparallel, base trisinuate; with narrow striae,

362 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

containing punctures distinct only near base; interstices each with a row of
distinct punctures. Abdomen with a shallow medio-basal depression. Femora
edentate. Length, 1:75-2:75 mm.

®. Differs in having the rostrum thinner, slightly longer than prothorax,
ridges finer, punctures smaller, antennae inserted somewhat nearer the middle,
and abdomen evenly convex.

Habitat.—North-western Australia: Fortescue River (W. D. Dodd).

The seventeen specimens obtained vary considerably in size, but only to a
slight extent in colour; the head and abdomen are sometimes infuscated, and the
sutural infuscation is more widely dilated towards the base on some specimens
than on others. It belongs to Section 1, and is structurally close to B. angophorae,
sororia, and bryanti, but those species are deep black. ,The only distinct spot
on the upper surface is the small medio-basal one on the pronotum, but there is
usually a very feeble spot near the middle on each side, on the elytra there
are usually a few scales on the odd interstices.

BaARIS ILLEPIDA, Nn. Sp.

Dull reddish-brown, scutellum, suture, metasternum and base of abdomen
deeply infuscated or black. Upper surface very sparsely clothed with whitish
seales, but a small medio-basal spot on pronotum and a less distinct one in middle
on each side; under parts moderately densely clothed.

Head with sharply defined punctures between eyes, becoming smaller and
shallower posteriorly. Rostrum moderately curved, slightly longer than, prothorax;
punctures at base as between eyes, becoming smailer in front. Antennae inserted
two-fifths from apex of rostrum. Prothorax almost as long as basal width, which
is fully twice that of apex; with rather dense, sharply defined punctures, becoming
smaller in front and larger on sides. Elytra slightly wider than prothorax at base,
sides nowhere parallel; with narrow striae, containing distinct punctures on basal
half; interstices each with a distinct row of transverse punctures, becoming larger
towards base, and crowded and irregular on second and third near base. Under
surface with punctures varying considerably in size and density. Femora edentate.
Length, 3-3-5 mm.

Habitat.—Northern Territory: Darwin (G. F. Hill).

Belongs to Section 1. Structurally and to a certain extent in colour and
clothing resembles the preceding species, but is larger and with a different
rostrum. B. bryanti, also from the Northern Territory, is deep black. The two
specimens taken appear to be females.

BARIS MELANOCHROA, Nl. SD.

dg. Black. Upper surface glabrous, under parts almost so.

Head with rather small but sharply defined punctures in front, elsewhere with
very minute ones. Rostrum the length of prothorax, strongly curved; with rather
dense, sharply defined punctures, becoming coarser and crowded on sides. Antennae
inserted about one-third from apex of rostrum. Prothorax longer than apical
width, which is only about half that of base; punctures not very close together
and rather small, but sharply defined, becoming larger and more crowded on Sides.
Elytra subcordate, outlines subcontinuous with those of prothorax; with narrow
striae containing deep-set punctures, but some fairly distinct ones towards base
and sides; interstices each with a row of scarcely visible punctures, but becoming
fairly distinct on sides. Most of sterna and tip of abdomen with crowded

BY A. M. LEA. 363

punctures, elsewhere (and especially on third and fourth segments) sparser and
smaller; first segment depressed along middle. Femora edentate. Length,
3-3-75 mm.

9. Differs in having smaller punctures between eyes, rostrum slightly longer
than prothorax, and abdomen evenly convex.

Habitat.—Queensland: Cairns district (H. Allen, F. P. Dodd and A. M. Lea).

Belongs to Section 1 of the genus, but is strikingly close in appearance to
B. cyaneotincta of Section 3. On several specimens the claw joint and parts of
the under surface are obscurely reddish. The specimen (a female) taken by
Mr. Allen has the punctures in front of the head somewhat larger than on the
other (8) specimens, and its rostrum is slightly gibbous at the base.

BaRIS VULNERATA, ND. Sp.

Black. Glabrous except for sparse white setae on under parts.

Head almost impunctate; with a large, round, deep, interocular cavity.
Rostrum rather wide and curved; with rows of distinct punctures, becoming
crowded and irregular on apex and sides. Antennae inserted two-fifths from apex
of rostrum. Prothorax almost as long as its basal width, sides obliquely narrowed
to apex; with sparse and very minute punctures, becoming fairly distinct about
apex, and denser and coarser on sides at base and apex. Elytra slightly wider
than prothorax, but their outlines almost continuous with those of prothorax;
with fine striae almost vanishing about middle of each elytron, and containing
punctures only near suture and sides. Metasternum with rather coarse, crowded
punctures; episterna arched downwards, each with a single row of punctures.
Basal segment with a shallow depression in male, apical segment with fairly large,
crowded punctures, sparse and small or absent elsewhere. Legs rather long,
femora edentate. Length, 2 mm.

Habitat—Queensland: Cairns district and Mount Tambourine (A. M. Lea);
Bellenden Ker (Dr. E. Mjoberg).

An elliptic species of Section 2, with elytral striae vanishing in parts; and
distinct from all other members of the subfamily by the large interocular fovea.
A specimen from Cairns has the prothorax and rostrum reddish, probably from
immaturity.

BARIS EPISTERNALIS, N. SDP.

Black. Upper surface almost glabrous; under parts with sparse white
setae, but each side piece of metasternum densely clothed.

Head minutely punctate, with a faint interocular impression. Rostrum long,
thin, and moderately curved; with rows of small punctures. Antennae inserted
about two-fifths from apex of rostrum. Prothorax decidedly transverse, apex not
half the width of base; with fairly dense, sharply defined punctures, becoming
crowded on sides. Elytra wide, slightly wider than prothorax, but with outlines
continuous with those of prothorax; with narrow striae, containing small
punctures; interstices each with a row of distinct punctures. Under surface with
irregularly distributed punctures, larger towards sides of metasternum than
elsewhere. Femora edentate. Length, 1:5 mm.

Habitat—Northern Queensland (Blackburn’s collection): Cairns district
(F. P. Dodd).

A minute rather wide species, with each metasternal episterna appearing as
a white line, although when greasy the white disappears. B. vagans is a

proportionately larger species, without special clothing on the metasternum.
T

364 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

B. australasiae, which has somewhat similar clothing on the metasternum, is a
smaller and decidedly narrower species, with seriate clothing on elytra. From
behind the base of the elytra appears bilobed, but from directly above it is seen
to be trisinuate. The male has a conspicuous depression on the basal segment
of abdomen. The pectoral canal is deep in front and bounded by carinated ridges,
it is quite as deep between the front coxae, but is narrower, so the species might
almost be considered as belonging to Solenobaris, but the subcontinuous outlines
of prothorax and elytra and the general appearance are as on many species of
Baris, to the second. section of which genus it seems desirable to refer it.

BARIS SETIPENNIS, D. Sp.

Black, antennae, tip of rostrum and parts of legs obscurely reddish. With
stramineous setae, on elytra forming a regular row on each interstice; on under
parts mostly white.

Head with small sharply defined punctures. Rostrum slightly longer than
prothorax, moderately curved, subgibbous at base, with dense punctures of
moderate size, becoming crowded at base and sides. Antennae inserted two-fifths
from apex of rostrum in male, almost in middle in female. Prothorax moderately
transverse, basal half almost parallel-sided, then strongly narrowed to apex; with
dense, sharply defined punctures of moderate size, becoming denser on sides.
Elytra slightly wider than prothorax, almost parallel-sided to beyond the middle,
base bilobed; with deep striae, containing deep-set punctures; interstices each
with a distinct row of punctures. Under surface with sharply defined, irregularly
distributed punctures, varying from small to fairly large. Femora edentate.
Length, 4:25-4-5 mm.

Habitat.—Queensland: Endeavour ‘River (Blackburn’s collection), Cairns
district (A. M. Lea).

The pectoral canal is rather short and narrow, but is bounded by well defined
ridges, and as the rostrum is subgibbous at base it seems best to refer it to
Section 3 of the genus, from all the previously described species of which it differs
considerably in appearance. It is allied to B. elliptica, but is larger and has
sparser clothing. It is about the size of B. basirostris, of Section 6, but has a
slight depression only instead of a deep notch between the head and base of
rostrum. In general appearance it is like several of the narrow species of
Melanterius of the Cryptorhynchides. The setae on the elytra are regularly
disposed, but about one-third of them are stouter than the others, and white. The
abdomen of the male is less convex than that of the female, and its rostrum is
slightly shorter, stouter, and with stronger punctures.

'

BARIS CYANEOTINCTA, ND. SD.

Black, prothorax and elytra with a metallic-blue gloss, becoming faint on
under surface; club and tip of rostrum obscurely reddish. Upper surface glabrous,
under parts with a small seta in each puncture.

Head with very minute punctures, and with a small interocular impression.
Rostrum about the length of prothorax, moderately curved, somewhat gibbous at
base; with numerous well defined but rather small punctures, becoming larger and
crowded on sides. Antennae inserted about one-third from apex of rostrum.
Prothorax slightly transverse, sides feebly diminishing in width to near apex, and
then strongly to apex itself; with sharply defined punctures of moderate size,
fairly numerous in middle, but crowded on sides. Elytra slightly wider than

BY A. M. LBA. 365

prothorax, nowhere quite parallel-sided, base trisinuate; with narrow striae
containing fairly distinct punctures; interstices each with a distinct row of small
punctures, becoming rather large about base. Sterna and apical segment of
abdomen with crowded and rather large punctures, smaller and sparser elsewhere.
Femora each with a small acute tooth. Length, 3-5-4 mm.

Habitat—Queensland (H. J. Carter), Cairns district (F. P. Dodd and
A. M. Lea).

A somewhat metallic species of Section 38, in general appearance, except
colour, like large B. sororia and angophorae, but prothoracic punctures less
crowded, and seriate ones on the interstices much less distinct. B. glabra is
smaller, with much smaller punctures, and more polished interstices. The male
differs from the female in having the rostrum slightly shorter, antennae inserted
slightly nearer the apex, and basal segment of abdomen with a slight depression.

BARIS NIVEODISPERSA, 0. SDP.

Black, shining, base of antennae and claw joints reddish. With depressed
white scales, forming small spots on sides at base and apex of prothorax, a distinct
transverse spot on each elytron at summit of apical slope, a few small spots or
single scales elsewhere, dense on metasternal episterna, and sparse on the rest
of the under surface, but legs and front of prosternum with some white setae.

Head with minute punctures. Rostrum slightly longer than prothorax,
strongly curved, subgibbous at base; with rows of punctures becoming smaller
and crowded in front, and coarse on sides. Antennae inserted about one-third
from apex of rostrum. Prothorax moderately transverse, sides gently rounded,
but suddenly becoming subtubular at apex; with sparse and small but sharply
defined punctures, altering to longitudinal scratches on sides. Elytra elongate-
subcordate, sides diminishing in width from near base; with narrow striae, in
which punctures are deeply set; interstices each with a row of very minute
punctures, becoming larger (but still small) at base, junction of fourth and fifth
on each elytron with a small tubercle immediately behind the white spot. Under
surface with irregularly distributed punctures of varying sizes, but two large
ones or small foveae in front of prosternum. Femora minutely dentate. Length,
4 mm.

Habitat—Queensland: Brisbane (H. W. Brown).

A well defined species of Section 4. The small elytral tubercles are rendered
more distinct by the surface behind them being depressed; B. bituberculata, with
somewhat similar tubercles, has very different punctures, and the upper surface
glabrous. The absence of elongated spots from the elytra distinguishes it from
the descriptions of B. albopicta and albigutta (the latter also belongs to Section 4).
B. niveonotata is somewhat more robust, with more numerous scales on elytra,
larger prothoracic punctures, and different rostrum. From B. apicinivea it differs
in the prothorax being without white vittae, and by the elytra having some white
scales on the basal third. The tips of the elytra are lengthened, so that the
pygidium is practically concealed. A second specimen differs from the type in
having the rostrum slightly longer and thinner, and with much less distinct
punctures on its upper surface, where also they are scarcely seriate; but its
abdomen is similar to that of the type.

BARIS PRAEMORSA, Nl. SD.
Black, parts of antennae obscurely diluted with red. Some snowy-white
seales at sides of base of rostrum, forming a small spot on each side in front of

366 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

prothorax (invisible from above), two elongate spots on third interstice: one at ~
base, the other beyond the middle, a spot on submarginal interstice, a spot on
each side of second segment of abdomen, and some on metasternum between the
coxae.

Head with rather dense and small, shallow punctures. Rostrum suddenly
elevated above head, fairly stout; with coarse, crowded punctures on sides and
base, becoming smaller in front. Antennae inserted one-third from apex of rostrum.
Prothorax almost as long as the basal width, sides feebly dilated to near apex
and then suddenly narrowed; with fairly dense, sharply defined punctures of
moderate size, becoming crowded and larger on sides. Elytra elongate-cordate,
base bilobed not much wider than prothorax; with narrow striae containing deep-
set punctures; interstices each with a row of small, irregular punctures, becoming
crowded posteriorly and larger towards base. Under surface with crowded
punctures, becoming smaller and sparser on parts of abdomen; basal segment of
abdomen of male with a shallow median depression; pectoral canal short and
feeble, but with two small foveae in front. Femora edentate. Length, 3-3-5 mm.

Habitat—Queensland: Cornwallis Island (C. T. McNamara), Magnetic Island
(A. M. Lea).

A well-marked species of Section 4; the spots noted are well defined and
constant on four specimens, but in addition a few single scales are scattered about.
It differs from the description of B. albiguita, with which it could be associated in
the table, in having only four spots (excluding the marginal ones) on the elytra.
Seen from the side there is a distinct notch between the head and base of rostrum.
The female differs from the male in having the rostrum slightly thinner, with the
punctures in front somewhat smaller and not quite so crowded, and the abdomen
evenly convex. One of the specimens from Cornwallis Island has the legs obscurely
diluted with red and the scape entirely red. The specimen from Magnetic Island
is of a piceous-brown, becoming black in parts.

BARIS LATERICOLLIS; Nn. Sp.

Black, shining. Upper surface glabrous, under parts sparsely clothed.

Head with small and fairly dense punctures. Rostrum slightly longer than
prothorax, strongly curved, slightly elevated above head at base, parallel-sided
to insertion of antennae (almost in middle) and then slightly narrowed to apex;
with numerous rather small, but sharply defined punctures on basal half, becoming
crowded and larger on sides, apical half with smaller and sparser punctures.
Prothorax moderately transverse, sides rounded, apex subtubular and not half
the width of base; with numerous small (but not crowded), sharply defined
punctures, becoming larger on sides, where many are connected by oblique
scratches. Elytra elongate-cordate, base not much wider than prothorax; with
narrow striae containing deep-set punctures, less distinct in middle than about
base and apex; interstices each with a series of minute punctures, becoming
irregular about base. Under surface with irregular punctures, crowded and fairly
large on parts of sterna, smaller and sparser on abdomen; pectoral canal almost
absent, but with two large punctures in front. Legs rather long, femora edentate.
Length, 4-4-5 mm.

Habitat.—Queensland: Cairns district (F. P. Dodd and Dr. E. W. Ferguson).

A large, polished species of Section 5; much larger than any glabrous species
previously known from Australia. In general appearance it is close to Myctides
balaninirostris, but the rostrum and prosternum are different. On each side, in

BY A. M. LEA. 367

front of the prothorax, the punctures for a short distance are connected by deep
striae, rather than scratches, so that from behind it appears to be almost notched
there. There are three fairly large but shallow depressions towards the side on
each elytron. One specimen has a shallow depression along the middle of the
two basal segments of abdomen, and as its rostrum has somewhat coarser
punctures than the other, and the antennae are stouter, it is probably a male,
the other probably being a female.

BARIS DILATATIFRONS, Nl. SD.

Black, somewhat shining. Upper surface glabrous, under surface and legs
very sparsely clothed.

Head with small, dense punctures; a shallow depression between eyes. Rostrum
about the length of prothorax, distinctly wider in front of insertion of antennae
(about two-fifths from apex), than behind same; with numerous but not very
sharply defined punctures, and with a rather feeble median ridge. Prothorax
about as long as its greatest width; with numerous, but not crowded, punctures
of fairly large size, becoming smaller and sparser in front. HEHlytra elongate-
cordate, near base about one-fourth wider than prothorax; narrowly striate, the
striae containing fairly large punctures towards base, these becoming rapidly
smaller towards middle, and disappearing posteriorly; interstices impunctate.
Under surface with punctures varying from rather sparse and small, on parts of
abdomen, to fairly dense and large on parts of sterna. Prosternum rather widely
grooved, the grooves bounded by a ridge on each side in front of the coxa.
Abdomen with a depression on basal segment, and another on apical segment.
Legs rather long and thin, femora scarcely perceptibly dentate. Length, 2-5-3 mm.

Habitat Queensland: Cairns district (F. P. Dodd).

An ordinary looking species, except that the rostrum is dilated in front,
so as to be suggestive of Cossonus. It is nearer Section 7 than any other of the
genus, but the pectoral canal is bounded by carinated ridges. The eyes are not
sufficiently coarsely faceted to entitle the species to be referred to Section 2. The
scutellum from some directions appears to be feebly bilobed. On one of the
three specimens, the middle coxae and parts of the antennae are obscurely reddish.

BARIS PULCHRIPENNIS, 0. Sp.

Black, most parts with a greenish gloss, elytra metallic blue, parts of legs
and tip of rostrum obscurely reddish. Elytra with elongated spots of white scales;
under surface and legs with irregularly distributed white scales, but becoming
dense on side pieces of metasternum.

Head with minute punctures, becoming fairly distinct in front. Rostrum
slightly longer than prothorax, moderately curved, rather thin; with dense and
rather small punctures, becoming larger on sides behind antennae. Prothorax
about as long as the greatest width (at extreme base), sides gently bisinuate
and strongly narrowed to apex; with sharply defined punctures, becoming larger
and more numerous about base than elsewhere. Elytra elongate-cordate, outlines
subcontinuous with those of prothorax, deeply and narrowly striate, the striae
containing punctures visible only from directly above; interstices with rows of
minute punctures, becoming fairly coarse about apex. Under surface with
punctures, varying from rather sparse and small on three median segments of
abdomen, to crowded and rather large. Prosternum feebly grooved in middle in
front. Legs rather long, femora edentate. Length, 4-4-5 mm.

Habitat—New South Wales: Clarence River (Blackburn’s collection).

368 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

This species cannot apparently be referred to any of the eight sections’
previously noted; the prosternum is shallowly grooved in front, but the groove
is not bounded by carinated ridges, and punctures or small foveae if present
are concealed by the clothing, which is denser only on the sides of the metasternum.
It apparently resembles B. albopicta, but the elytral spots are somewhat differently
disposed, and the femora are edentate. The prothorax is more triangularly shaped
than in B. niveonotata, and its punctures are smaller. There are two distinct and
two feeble spots on each elytron; of the distinct spots the first is an elongated
one on the third interstice at the base, and the second is on the fifth interstice
at the apical third; the small spots (apparently easily abraded) are on the fifth
interstice near the base, and on the third near apex. On parts of the third and
fifth interstices, there are two quite regular rows of punctures, on each of the
others there is but one row, but towards the sides they become irregular and
fairly numerous. On three specimens the rostrum is metallic green almost to the
tip, on a fourth it is obscurely reddish throughout; on one the suture is of a
different blue to the rest of the elytra, but this is probably a post mortem variation.

BARIS ANGOPHORAE Lea.
Trans. Roy. Soc. 8S. Aust., 1906, xxx, p. 86.

Five specimens from Stradbroke Island and Bunya Mountains probably belong
to this species, but each of them has a row of sparse white depressed setae in the

alternate interstices of elytra; on the types there are fewer similar setae, but
others may have been abraded.

BARIS TRANSVERSICOLLIS Lea.
Proc. Linn. Soc. N.S.W., 1914, xxxix, p. 686.
The types were evidently partly abraded. Numerous specimens from Cairns

are in perfect condition, and have a distinct row of scales sometimes semidouble,
on each elytral interstice.

BARIS SORORIA Lea.
Trans. Roy. Soc. S. Aust., 1906, xxx, p. 87.

Numerous specimens of this species from Queensland are larger than the type
(almost up to 4 mm.). Fresh localities are Brisbane, Cowell Creek, and Magnetic,
Thursday, and Moa Islands.

BARIS MICROScCOPICA Lea.
Trans. Roy. Soc. 8. Aust., 1906, xxx, p. 89.

A specimen from Cairns agrees with the types of this species, previously
known only from Western Australia.

BARIS GERALDTONENSIS, new name.
B. oblonga Lea, Trans. Roy. Soc. S. Aust., 1906, xxx, p. 89.

Baris oblonga having been used by Bohemann in 1859 for an ex-Australian
species, I have to propose the above substitute for the Australian species.

BARIS CAIRNSENSIS, new name.
B. glabra Lea, Trans. Roy. Soc. S. Aust., 1906, xxx, p. 91.

Baris glabra was used by Herbst in 1784.

BY A. M. LBA. 369

BARIS BARRONENSIS, new name.

B. ebenina Lea, Trans. Roy. Soc. S. Aust., 1906, xxx, p. 95.
Baris ebenina was used by Pascoe in 1885.

IpSICHORA TIBIALIS, 0. Sp.

Black, most parts with a slight greenish gloss, elytra deep blue. Upper
surface glabrous, under surface and legs with short, sparse clothing.

Rostrum moderately curved, about one-fourth shorter than prothorax, upper
surface shining and with sparse and small punctures about base, becoming
numerous at apex, on the sides dense and rather coarse. Prothorax distinctly |
shorter than the basal width, which is about twice that of the apex; with
numerous but not crowded punctures of moderate size, becoming somewhat denser
on sides. Elytra cordate, outlines continuous with those of prothorax; with
regular rows of round punctures, the interstices with very minute punctures. Under
surface with punctures varying from small and sparse on parts of abdomen, to
dense and fairly large on parts of sterna. Basal segment of abdomen with a
shallow median depression, apical segment with a round one. Femora edentate,
front tibiae with a small downward projection at apical third. Tength, 3-75 mm.

Habitat—Queensland: Cairns (H. Allen). Unique.

As its femora are edentate this species in the table of Ipsichora could be
associated with J. desiderabilis, from which it differs in being less brightly
coloured, front tibiae with a small subapical tooth (closer to the apex than on
Baris hoplocnemis) and apical hook smaller and black. On the type the head
has been forced back so as to conceal most of its base. The larger punctures on
the elytra are in regular rows, but not in striae, except that about the suture
and sides (but not to base) a few striae are present. The type appears to be a male.

IPSICHORA BIMACULIBASIS, ND. SD.

Brilliant metallic blue, altering to green on head and rostrum, under surface
and legs partly blue and green; parts of antennae and of tarsi obscurely reddish.
Elytra with an elongated spot of white scales on third interstice at base, under
surface, and under surface of femora, moderately clothed with white scales.

Head with small punctures, sharply defined only near eyes. Rostrum almost
as long as prothorax, moderately curved; with numerous small, sharply defined
punctures, becoming larger and more crowded on sides. Prothorax about as long
as the basal width, sides feebly bisinuate and strongly narrowed to apex; with
fairly numerous and small, but sharply defined punctures. Elytra elongate-
cordate, outlines subcontinuous with those of prothorax; narrowly and deeply
striate, the striae containing punctures visible only from directly above. Under
surface with punctures varying in size and density with their positions. Legs
rather long and thin, femora edentate. Length, 3 mm.

Habitat Queensland: Cairns (F. P. Dodd). Unique.

Smaller than the previously named Australian species of the genus and
elytra with two white basal spots. The front of the prosternum is similarly
clothed and in general is close to that of Baris pulchripennis, but as it is a
conspicuously blue species it appears better to refer it to Ipsichora than to a
doubtful section of Baris. The two basal segments of the abdomen have a large
shallow depression, so that although the apical segment is non-foveate, the type
is probably a male.

tea, Trans. Roy. Soc. 8S. Aust., 1906, xxx, p. 98.

370 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

IPSICHORA DESIDERABILIS Lea.
Trans. Roy. Soc. 8. Aust., 1906, xxx, p. 98.

Two males of this species are unusually small (3 mm.). Several have the
pronotum more purplish than blue. On greasy specimens the metallic gloss is lost,
and the derm becomes black.

SOLENOBARIS CRYPTORHYNCHOIDES, 0. SP.

6. Black, highly polished. Under surface and legs with sparse, short clothing.

Head with dense and small but well defined punctures. Rostrum almost the
length of prothorax, moderately curved and stout, dilated in front of antennae
(these inserted slightly in front of middle); upper surface with small punctures,
becoming crowded in front, and rather coarse on sides. Prothorax almost as long
as the basal width, which is much wider than apex; punctures sparse and small
in middle, becoming denser about base and apex, and denser and fairly large on
sides. Elytra cordate, outlines continuous with those of prothorax; with rows of
fairly large punctures, rapidly becoming smaller and disappearing beyond the
middle; narrow striae near suture and sides only. Punctures small and sparse
on parts of abdomen, larger and denser elsewhere; basal segment with a shallow
depression continued on to metasternum, apical segment with a shallow sub-
triangular one. Legs long, femora acutely dentate. Length, 2-5-2:75 mm.

©. Differs in having the rostrum somewhat longer and thinner, with smaller
punctures, antennae inserted slightly nearer base than apex, abdomen without
depressions, and legs somewhat shorter.

Habitat— Queensland: Cairns district (F. P. Dodd and A. M. Lea).

At first glance the pectoral canal appears to terminate against a narrow
transverse wall belonging to the mesosternum, but on separating the prothorax
from the mesothorax the wall is seen to be part of the former; on this account,
and also the continuous outlines of prothorax and elytra, the species should
perhaps have been referred to a new genus; it is certainly very different from
the other species of Solenobaris. With the rostrum at rest in the canal the
resemblance to Idotasia of the Cryptorhynchides is a striking one; but the distinct
pygidium and side pieces of mesosternum thrust like a wedge between the elytra
and prothorax, are typical of the Baridiides.

SOLENOBARIS EDENTATA Lea.
Trans. Roy. Soc. S. Aust., 1906, xxx, p. 103.

Three specimens, from Cairns, differ from the types in being slightly
larger (2-2-5 mm.) and with a faint bluish gloss on the elytra. The male has a
shallow depression on the first segment of abdomen, and another on the fifth,
and its eyes are even closer together than on the female. In both sexes the
rostrum is slightly dilated at the insertion of antennae, but in the female it is
subulate in front of them.

LEPIDOBARIS, Nn. &g.

Head moderately large. Eyes large, ovate, obtusely pointed below, widely
separated, finely faceted. Rostrum long, thin, and strongly curved. Antennae
inserted in middle of rostrum in male, nearer the base in female; scape not quite
extending to eyes. Prosternum faintly depressed along middle in front. Without
visible pygidium. Front coxae widely separated; femora edentate. Body densely
sguamose. Other characters as in Baris.

BY A. M. LEA. dial

The only known species appears at first to belong to Baris, but there is no
visibie pygidium; the prosternum is faintly depressed along the middle but without
ridges there, as in Gymnobaris; from that genus also it differs in its clothing,
edentate femora, insertion of antennae, etc. Genotype L. metasternalis, un. sp.

LEPIDOBARIS METASTERNALIS, Nl. Sp.

6. Dark reddish-brown, head almost black, antennae legs and part of rostrum
reddish. Densely clothed with variegated scales on upper surface, white on under
surface and legs, head and rostrum sparsely clothed.

Head subopaque (faintly shagreened) with numerous small punctures. Rostrum
slightly longer than prothorax; with numerous small, sharply defined punctures,
becoming rather coarse on sides behind antennae. Antennae inserted almost in
exact middle of sides of rostrum, scape much shorter than funicle, its tip distant
from eyes about two-thirds the length of the following joint. Prothorax moderately
transverse, basal half subparallel, then strongly rounded to apex, base bisinuate;
with crowded, partially concealed punctures, and with a feeble median line. HElytra
subcordate, outlines subcontinuous with those of prothorax, widest near base;
narrowly and deeply striate, the striae with deep-set punctures; interstices with
crowded, mostly concealed punctures. Basal segment of abdomen slightly depressed
in middle. Length, 3:5-3-75 mm.

9. Differs in being wider, rostrum thinner, conspicuously longer than prothorax,
with sparser clothing, and smaller punctures on sides behind antennae, these
inserted nearer the base, with the tip of the scape nearer the eyes, prothorax
more transverse, its sides more evenly rounded, and abdomen larger and more
strongly and evenly convex.

Habitat—New South Wales: Eccleston, in flood debris (J. Hopson).

The head is clothed only near eyes, the rostrum only on the sides behind the
antennae. On two females the clothing on the prothorax and elytra is mostly
mouse-coloured, with scattered whitish scales on the former; on the latter there
is a white vitta on the suture from the base to beyond the middle, where it is
joined to an oblique vitta on each side on to the fourth interstice, single whitish
seales are also scattered about; on the under surface the clothing is dense, notably
on the metasternal episterna; just before each episternum there is a narrow shining
line, with a row of squamiferous punctures. On the male the clothing on the
upper surface is paler and less variegated, but the sutural and oblique vittae are
distinct; on the sides of its prothorax some of the scales have a slight golden
gloss. The elytra are really flattened along the middle of the apical slope, but
from some directions, owing to the adjacent clothing, they appear to be slightly
concave there..-

APINOCIS, 0. &.

Eyes widely separated and with facets of moderate size. Rostrum long, thin,
and strongly curved. Prosternum not at all concave along middle. Pygidium
small. Front coxae widely separated. Femora edentate. Other characters as in
Baris.

The prosternum is entirely without a depression along the middle, the surface
there being in fact rather strongly convex; this precludes the species from being
referred to Baris. In the fifth section of that genus all the species are without
a longitudinal groove, but there are two distinct punctures or small foveae in the
transverse subapical constriction; on A. variipennis the constriction is deep, but

372 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

is entirely without foveae. The pygidium is small, and on one specimen is°
concealed, but evidently from pressure. Genotype A. variipennis, n. sp.

APINOCIS VARIIPENNIS, 0. Sp.

¢. Reddish-castaneous, some parts. darker. Irregularly clothed with
stramineous setae or thin scales, becoming paler on under surface and legs.

Head very minutely punctate. Rostrum slightly longer than prothorax,
feebly dilated at base and apex; with small punctures, seriate in disposition behind
antennae, irregular in front, and larger on the sides than elsewhere. Antennae
inserted about two-fifths from apex of rostrum, scape not quite extending to eyes.
Prothorax almost as long as the basal width, which is about twice that of apex;
with irregularly distributed punctures of various sizes, but mostly small. Elytra
elongate-cordate, sides subcontinuous with those of prothorax; with narrow striae.,,.
containing fairly large punctures about base, but becoming small posteriorly;
interstices each with a row of punctures, fairly large about base, very small
elsewhere. Under surface with larger and denser punctures than on upper surface.
Two basal segments of abdomen with a large, shallow depression. Length,
3-5-3-75 mm.

9. Differs in having the rostrum distinctly longer and thinner, antennae
inserted nearer the base than apex of rostrum, prothorax slightly shorter,
and abdomen more strongly and evenly convex.

Habitat.—Victoria: Natya and Cheltenham in October, Coburg in June (C.
Oke), Melbourne (National Museum). Type in South Australian Museum.

The four specimens under examination all vary in colour; the only male has
the head, pronotum and metasternum darker than the adjacent parts, but not
black, the elytra have the base and apex paler than the median parts, but the two
shades not sharply limited. All the females have the mesosternum, metasternum,
and abdomen black or blackish; on the Natya specimen the upper surface is entirely
reddish, on the one from Cheltenham it is somewhat as on the male, except that
the median part of the elytra is almost black, with two rather conspicuous pale
spots behind it; on the Melbourne specimen the upper surface is black, except
that all the margins of the prothorax, and the base and apex of elytra are
obscurely reddish. The clothing on the Cheltenham specimen is fairly dense on
the margins and middle of pronotum, and about base and apex of elytra, it also
forms a single distinct row on each interstice; from the others it has evidently
been partly abraded; as it is sparser on the prothorax and elytra, and on the
latter the rows are absent from the interstices. On the under surface the setae
are moderately and almost evenly distributed. The largest punctures on the
pronotum are about as large as the largest on the elytra and are irregularly placed
on each side of the middle, the middle itself being almost impunctate.

PENTARTHRUM.

In the revision of the Cossonides by Wollaston’, Pentarthrum is referred to on
pp. 434, 446, 513 and 598-601; but he nowhere notes the separation of the front
coxae. He states, however, that the type of the genus is the British P. huttoni,
and on that species the front coxae are widely separated; on the New Zealand
P. zealandicum the front coxae are also widely separated. Broun,? when revising
the New Zealand species of the genus Pentarthrum and proposing many new genera

1 Wollaston, Trans. Hunt. Soc. Lond., 1873.
4Broun, Trans. N.Z. Inst., 1908, xli, p. 155.

BY A. M. LEA. 373

at its expense, leaves in the genus those species having “coxae widely separated,
the anterior included’. Of the named Australian species at present standing in
the genus, P. nigrum (if the specimens before me are correctly identified) has the
front coxae rather feebly separated, in P. millingtoni the separation is about
the same, in P. nepeanianum the separation is wide, in P. awstrale they are fairly
widely separated, although not as widely as in huttoni or zealandicum. Wollaston,
however, when describing a species from Japan, says of the legs, “antici ad basin
parum . . . distantes’’.t Of the following species only the first, P. orthodoxum,
will probably remain in the genus, as all the others have the front coxae more or
less close.

PENTARTHRUM ORTHODOXUM, D0. Sp.

6. Blackish-brown, legs and antennae paler.

Head with dense and sharply defined but not very large punctures in front,
disappearing posteriorly; with a slight interocular impression, continued, but very
feebly, almost to apex of rostrum. Rostrum flattened and straight, about as long
as the width at base of prothorax, about twice as iong as its own apical width,
parallel-sided to near base, where it is triangularly dilated and with punctures as
between eyes, smaller punctures elsewhere. Antennae inserted in middle of
rostrum, first joint of funicle longer than second, second than third, and third
than fourth, fourth and fifth transverse. Eyes small and prominent. Prothorax
somewhat pear-shaped, widest about one-fourth from base, constricted near apex,
where the width is but little more than half of that near base; with dense but
not very large punctures, becoming crowded on sides. Elytra parallel-sided to
near apex, slightly narrower than greatest width of prothorax; striate-punctate, -
punctures large and wider than interstices, each of these with an inconspicuous
row of punctures. Under surface with dense punctures, much smaller than on
pronotum, but becoming almost as large on sides; front coxae widely separated;
basal segment of abdomen with a wide, shallow, median depression, apical
segment with a shallow semicircular one. Length (excluding rostrum), 3-3:25 mm.

®. Differs in having the rostrum thinner and quite cylindrical almost to
extreme base, with smaller punctures and without median line, abdomen without
depressions and legs somewhat thinner.

Habitat—Tasmania: Hobart, Frankford (A. M. Lea), Franklin, Launceston
(British Museum).

With the rostrum sexually variable as in P. zealandicum, but a smaller
species and differently coloured; the punctures of the under surface are much
smaller than on P. nigrum. The knees are somewhat darker than the rest of the
legs; the club appears paler than the rest of the antennae, but this is due to
minute whitish pubescence. The longitudinal impression on the rostrum of the
male is very feeble and usually traceable from oblique directions only. The
pronotum has a faint medio-basal depression, which in some lights is traceable
almost to the middle. The distance between the front coxae is about equal to the
width of the apex of rostrum.

PENTARTHRUM FOVELVENTRE, 0D. SD.
6. Dark brown, antennae and legs slightly paler.
Head with numerous (but not crowded) sharply defined punctures of rather
small size in front, vanishing posteriorly. Rostrum subcylindrical, slightly

1 Wollaston, Trans. Ent. Soc. Lond., 1873, p. 14.

374 NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

curved, almost twice as long as the basal width; basal half with punctures about
as large as on front of head, but more crowded, smaller and sparser on front half.
Antennae inserted slightly nearer apex than base of rostrum. Eyes small and
prominent. Prothorax slightly constricted near apex, sides obliquely dilated to
near base, where the greatest width is about once and one-half of that of apex;
punctures rather dense (but not crowded) and of moderate size. Hlytra parallel-
sided to near apex, the width of widest part of prothorax; striate-punctate,
punctures large and mostly wider than interstices, each of these with an
inconspicuous row of punctures. Under surface with coarser punctures than on
pronotum but somewhat irregular; front coxae slightly separated; metasternum
and two basal segments of abdomen with a wide shallow depression, apical
segment with a large round fovea. Length, 3-3-5 mm.

9. Differs in having a somewhat smaller rostrum, without a depression
common to metasternum and abdomen and apical segment not foveate.

Habitat—Tasmania (Aug. Simson): Franklin (British Museum), Hobart
(A. M. Lea).

At first glance strikingly close to the preceding species (the British Museum
specimen was, in fact, mounted on a card with a male of that species), but with
front coxae much less widely separated (almost touching), rostrum slightly
curved, not grooved along middle, and apical segment of abdomen of male
foveate. The club is clothed with minute white pubescence, in consequence of
which it appears paler than the rest of the antennae.

PENTARTHRUM FOVEICEPS, 0. Sp.

6. Black, antennae and legs obscurely diluted with red.

Head with a small interocular fovea, and with sharply defined and fairly
dense, but not crowded, punctures of moderate size, terminated before base.
Rostrum slightly curved, about as long as greatest width of prothorax, sides
gently incurved to middle; punctures at base much as on front of head, becoming
smaller and more numerous in front. Antennae inserted in middle of rostrum;
first joint of funicle as long as second and third combined, second to fifth
transverse. Eyes small’ and prominent. Prothorax about twice as long as the
apical width, sides slightly constricted near apex, then slightly dilated posteriorly,
and rounded to base itself, which is slightly wider than apex; punctures dense,
sharply defined and fairly large, becoming crowded on sides. Elytra parallel-
sided to near apex, striate-punctate, punctures large and wider than interstices,
each of these with a row of minute punctures. Prosternum with crowded
punctures, many of which are transverse; front coxae slightly separated; basal
segment of abdomen with a shallow depression, apical segment with a semi-
circular one. Length, 4-5 mm.

2. Differs in being somewhat smaller, rostrum thinner, abdomen simple, and
legs shorter.

Habitat—Tasmania: Hobart, Frankford (A. M. Lea), Franklin (British
Museum).

About the size of P. nigrum, but prothorax constricted near apex and with
somewhat smaller punctures; on the female the rostrum is narrower towards the
apex than on the male, but it is not truly cylindrical as on P. orthodoxum and
P. zealandicum. The funicle is somewhat darker than the rest of the antennae;
on most specimens the base of the head is obscurely diluted with red. On several
there are remnants of an impunctate median line on the pronotum.

Se

BY A. M. LEA. 375

PENTARTHRUM INTEROCULARE, 0D. SD.

6. Black, antennae and legs obscurely diluted with red.

Head with sharply defined and numerous, but not crowded, punctures of
moderate size in front; without an interocular fovea. Rostrum about as long
as width of apex of prothorax, sides feebly diminishing in width to antennae,
thence parallel-sided to apex; with punctures at base and between eyes, becoming
smaller and more numerous to apex. Prothorax somewhat pear-shaped, slightly
constricted near apex, dilated to near base; with rather dense, sharply defined
and fairly large punctures, becoming crowded and longitudinally confluent on
sides. Elytra parallel-sided to near apex; striate-punctate. Prosternum with
punctures much as on middle of pronotum; basal segment of abdomen with a
shallow median depression, the apical segment with a semicircular one. Length,
3-3-5 mm.

Q. Differs in having a slightly thinner rostrum and basal segment of abdomen
evenly convex.

Habitat—Tasmania (Aug. Simson): Franklin (British Museum), Hobart
(C. E. Cole), Launceston (A. M. Lea).

Somewhat smaller than the species I have identified as P. nigrum, and with
more crowded punctures on the sides of prothorax, many being confluent. In many
respects it is close to P. foveiceps, but is smaller, head without an interocular
fovea, rostrum parallel-sided in front of antennae, prothorax wider, with coarser
punctures, mostly longitudinally confluent on the sides, and prosternum with
sparser, larger and rounded punctures.

PENTARTHRUM STRIGICOLLE, N. sp.

dg. Black or blackish, parts of antennae and legs obscurely diluted with red.

Head with numerous well-defined punctures of moderate size in front, but
absent from base; with a small interocular impression, continued as a feeble
groove on basal half of rostrum. Rostrum about as long as width of base of
prothorax, slightly curved, sides feebly incurved to middle, with minute punctures,
somewhat larger at base. Antennae inserted almost in middle of rostrum. Eyes
small and prominent. Prothorax twice as long as the basal width, sides
constricted near apex, dilated posteriorly and evenly rounded at base; with
sharply defined and numerous comparatively small punctures, becoming larger on
sides but not confluent. Elytra parallel-sided to near apex, striate punctate.
Prosternum densely punctate, becoming strigose at apex and near coxae; front
coxae almost touching; basal segment of abdomen with a wide shallow depression,
apical segment semicircularly impressed. Length, 4:5-5 mm.

®. Differs in having the rostrum thinner and without a median groove, and
basal segment of abdomen evenly convex.

Habitat—New South Wales. Illawarra (H. W. Brown), Sydney, in September
(British Museum from J. J. Walker). ;

Near P. foveiceps, but rostrum of male feebly grooved, prothorax with smaller
punctures, prosternum transversely strigose and abdominal depression deeper
in the middle. On several specimens the sterna, and on one, the elytra, are
dark brown instead of black; on most of them the antennae, except the club,
are almost as dark as the rostrum.

NoOTIOSOMUS INSULARIS, Nl. SD.
6. Deep black and shining, antennae and tarsi obscurely reddish; club paler.
Head dilated and almost imptnctate about base, with distinct punctures and

376 ‘ NEW SPECIES OF AUSTRALIAN COLEOPTERA, Xix,

a small fovea between eyes. Rostrum about once and one-half as. long as apical
width, slightly curved, parallel-sided in front of antennae, incurved at their
positions, punctures slightly coarser and more crowded than those between eyes.
Antennae inserted slightly nearer base than apex of rostrum, first joint of
funicle about as long as second and third combined, second to seventh strongly
transverse and subequal. Prothorax about twice as long as the apical width,
constricted near apex, sides dilated to near base, which is about one-fourth
wider than apex and finely margined; with sharply defined and fairly dense
punctures, about as large as those between eyes, becoming crowded on sides;
with a feeble, shining median line. Scutellum small. Elytra parallel-sided to
beyond the middle, as wide as widest part of prothorax; striate-punctate, punctures
large, becoming smaller posteriorly; interstices with minute punctures and in
places faintly wrinkled. Under surface with sharply defined punctures, larger
and denser on prosternum and sides of metasternum than elsewhere. Basal
segment of abdomen with a wide shallow depression, slightly continued on to
second, sutures between second to fifth segments wide, and each with a row of
large punctures. Length, 4-5 mm.

9%. Differs in having the rostrum slightly shorter less incurved at insertion
of antennae and abdomen evenly convex.

Habitat.—Norfolk Island (J. C. Wiburd and A. M. Lea), Lord Howe Island
(A. M. Lea and wife).

This species is stouter and less depressed than the mainland forms of
Notiosomus, and the rostrum is nearer to that of Cossonus, the third tarsal joint
is narrow and entire. It is still more robust and with a much wider (but not
shorter) rostrum than the three New Zealand species of Phlaeophagosoma known
to me (P. corvinum, P. dilutum and P. pedatum). The rostrum is too long for
Stereoborus. The front tibiae are feebly bisinuate on the lower surface, the
apical sinus with fine pubescence invisible from several directions. The specimens
from Lord Howe Island have slightly coarser punctures and more wrinkled
elytra than the others, but the differences are very slight. Abundant on both
islands.

ISOTROGUS IRREGULARIS, N. SDP.
Black, antennae and legs more or less obscurely reddish.

Head with a distinct interocular impression, continued as a groove to
middle of rostrum, with a few small punctures near eyes. Rostrum slightly
curved, very little longer than wide, sides slightly incurved at insertion of
antennae (somewhat nearer apex than base) and but little dilated in front;
punctures denser and more distinct than on head, but still small. First joint of
funicle fairly long, the second shorter, third to seventh strongly transverse.
Prothorax more than twice as long as the apical width, sides dilated posteriorly
and rounded off at base, which is considerably wider than apex; with irregular
and rather large punctures on each side of a narrow median line, somewhat
similar punctures on sides, the intervening spaces with minute ones. Elytra
parallel-sided to near apex, almost the exact width of widest part of prothorax;
strongly striate-punctate, the interstices impunctate. Under surface with ‘large
and irregular punctures on sides, on prosternum, forming a semidouble row
between middle coxae, in middle of basal segment of abdomen on apical segment
and in some of the sutures. Length, 3-5-4 mm.

BY A. M. LEA. 377

Habitat.—Northern Territory: Darwin (G. F. Hill and British Museum).

A depressed species. The conspicuous punctures on each side of the median
line of the pronotum, although not in single rows, indicate that it is allied to
Isotrogus bilineatus and I. castelnaui, and it is therefore referred to Isotrogus,
although as Pascoe wrote when dealing with the former species, “The genus
might well be united to Cossonus”. From bilineatus it differs in having the rostrum
decidedly shorter, straighter, and with a longitudinal groove, as well as in the
prothoracic punctures. From castelnawi (which has more irregular rows of
punctures than on bilineatus, but still in single series) it differs in being smaller,
and with a conspicuous groove on the rostrum, in continuation of the interocular
impression. The male has a shallow depression on the basal segment of abdomen,
the female being flattened there, these being apparently the only external
indications of sex. The funicle is slightly darker than the rest of the antennae;
the legs are almost black.

Ten specimens, from Roebuck Bay, in the British Museum, appear to
represent a variety; they differ in being of a rather dark brown, with the
elytra bright castaneous, except that the suture and tips are slightly infuscated,
parts of the under surface and of the legs are also castaneous.

ADDITIONAL FLORA OF THE COMBOYNE PLATEAU, 1926.

By E. C. CHisHotm, M.B., Ch.M.
[Read 29th June, 1927.]

In These ProcrEpines, 1925, p. 284, I gave an account of the general
conformation and flora of the Comboyne Plateau; I now present some additional
records of the plants found there.

Fungi: Polyporus (Ovinus) mylittae Cooke.—A stipitate fungus of the family
Polyporaceae the sclerotium of which is frequently ploughed or dug up on the
basalt, the pileus or fructifying portion being rarely seen and apparently being
thrown above ground only when conditions are favourable. A specimen of
sclerotium brought to me was dark brown in colour and of hard consistency,
roughly oval in shape and concavo-convex on its broadest surfaces, measuring
six inches by three and a half inches and weighing ten ounces. Its interior on
fracture is of a creamy white colour. This is known as “Blackfellow’s bread’.

Muscineae: Dawsonia superba—This moss is fairly common, growing in
patches on mixed strata, especially in shady and moist gullies.

Polypodiaceae: Arthropteris obliterata R.Br.—A climbing fern found on
stems of trees; only seen on the eastern side of the Plateau growing on mixed
strata.

Cyclophorus serpens Forst.—This and C. confluens are mostly found on basalt
covering the trunks of Sambucus xanthocarpa (Native Elderberry), for which
they seem to have an attraction.

Platycerium grande A. Cunn.—An epiphyte on trunks and branches of brush
trees. This is quite a rare form here though quite common in the brushes near
sea level. It is only quite lately that I have. come across it on the Plateau.

Typhaceae: Typha angustifolia Linn.—Bullrush; plentiful, growing in the
creeks.

Liliaceae: Philydrum lanuginosum Banks.—An aquatic plant not very
plentiful, only seen on the eastern side of the Plateau. It has a conspicuous
yellow flower.

Kreyssigia multiflora Reichb.—A low growing plant with a pale mauve flower.
Only found on the eastern side of the Plateau growing on mixed strata.

Cordyline stricta Endl.—Palm Lily, not common and only seen on the eastern
side. It bears a handsome mauve flower.

Orchidaceae: Dipodium punctatum R.Br.—Uncommon and only found off the
basalt on the mountain side.

Microtis porrifolia R.Br.—A small orchid with an inconspicuous greenish
flower. It grows fairly plentifully on the cleared land.

Moraceae: Ficus henneana Mig.—A rare fig here, its habitat appears to be
further north. I know of only one tree.

Ficus eugenioides F.v.M.—Common in the brushes, having small leaves and
small yellow fruit.

BY E. C. CHISHOLM. 379

Proteaceae: Persoonia linearis Andr.—Not common and found on sedimentary
soil.

Loranthaceae: Loranthus pendulus Sieb—The commonest mistletoe of the
hardwood country and found mostly on the fringe of the Plateau growing on
Eucalypts.

Lauraceae: Cryptocarya sp.—A tall tree with blue fruit commonly called
“Corduroy Beech” from the fact that its trunk has a twisted appearance and is
deeply fluted. This is fairly plentiful in the brushes.

Pittosporaceae: Pittosporum revolutum Ait.—Not very plentiful. It has a
yellow flower and is found in the brushes.

Leguminosae: Oxylobium trilobatum Benth.—I have met with this only on
the eastern side growing on mixed strata.

Goodia lotifolia Salisb—Not plentiful and found mostly on the northern side
off the basalt. It bears a yellow flower.

Swainsonia coronillifolia Salisb—This grows in the same situation as the
last and is frequently seen just off the edge and down the mountain side. It has
a mauve or light purple flower.

Tremandraceae: Tetratheca thymifolia Sm.—Only found growing on the
eastern side on mixed strata.

Sapindaceae: Guioa semiglauca Radlk.—Not plentiful and found growing on
basalt.

Violaceae: Viola betonicifolia Sm.—Uncommon and only found on the out-
skirts of the Plateau amongst hardwood timber.

Myrtaceae: Eugenia australis Wendl—Rare, found growing some distance
down the southern side of the Plateau on basalt, having large elongate crimson
fruit and a white flower. It is one of the “Brush Cherries”.

Eugenia coolminiana C. Moore.—Another “Brush Cherry”, the fruit being
pink in colour, of large size and flattened at the poles. It is an inhabitant of the
brushes and grows into a fine tree.

Labiatae: Brunella vulgaris A. DC.—Uncommon, found growing on basalt.

Campanulaceae: Lobelia trigonocaulis F.v.M.—Plentiful, growing on the basalt
amongst the brush timber.

Wahlenbergia gracilis A.DC.—Bluebell, not very plentiful, found on basalt
but of stunted growth, the flowers being very small.

Goodeniaceae: Goodenia sp.—Apparently a new species, found on mixed strata
on the eastern side of.the Plateau. The affinities of this are being worked out
at the National Herbarium, Botanic Gardens.

Compositae: Gnaphalium purpureum L.—Common on cleared land on the
basalt.

ON THE PHYLOGENY OF SOME DIPTERA BRACHYCERA.
By G. H. Harpy,

Walter and Eliza Hall Fellow in Economic Biology,
Queensland University, Brisbane.

(Two Text-figures.)

[Read 29th June, 1927.]

The section of Diptera referred to here has long been known as the
Brachycera division of the suborder Orthorrhapha, but Dr. R. J. Tillyard (1926,
p. 345) has reversed the status of these two names making Brachycera a suborder
that incorporated all Diptera except the Nematocera and the term Orthorrhapha
becomes the name of the division.

The relationship between the various families is deduced from the researches
of various authors. The families are fast becoming better understood so it is
possible to illustrate their supposed affinities diagrammatically and to discuss
them from this aspect. J. R. Malloch (1917, pp. 173-181) dealt with the larval
and pupal characters of these families and he formed six superfamilies which
are now reduced by Tillyard to three as instanced by the table given below.
The simplicity brought about by this reduction has much to commend it.

Superfamily name used by:
Family.
Malloch. Tillyard.
LEPTIDAE TABANOIDEA
TABANIDAE
STRATIOMYIIDAE
(Some non-Australian STRATIOMYIOIDEA TABANOIDEA
families)
NEMESTRINIDAE } CYRTOIDEA
CYRTIDAE
SCENOPINIDAE THEREVOIDEA
THEREVIDAE
ASILIDAE ASILOIDEA
APIOCERIDAE ASILOIDEA
MYDAIDAE
BoOMBYLIIDAE
EXMPIDIDAE EMPIDOIDEA EMPIDOIDEA
DOLICHOPODIDAE

BY G. H. HARDY. 381

A diagram of the relationships between these families is very readily formed
and it is the purpose of this paper to point out some of the more striking features
arising from this method of allying the families, to indicate some of the weaker
points and to incorporate some of the lines along which my own researches are
tending to show a development regarding the superfamily Asiloidea.

The genealogical tree given below contains three main branches, the first
of these is the superfamily Tabanoidea into which come all forms containing
a pulvilliform empodium, and it is further divided into three divisions, the
Tabanid-Leptid Stem, the Stratiomyiid stem and the Cyrtid-Nemestrinid stem.

The first two of these contains insects that are largely associated with water,
many breeding in marshes and such-like places, but the third, the Cyrtid-
Nemestrinid stem is not associated in this manner. It is of interest to note
that the land-breeding family Nemestrinidae contains species that have a wing-
venation in which there is a marked tendency for the apex of the veins to move
towards the costa and this is repeated in the other entirely land-breeding section
the Asiloidea. In the light of our present knowledge this must be regarded as
parallel development but possibly it has a deeper significance, for the position of
the Cyrtid-Nemestrinid stem within the superfamily Tabanoidea appears to be
somewhat anomalous, differing so markedly as it does in many structures. It has
apparently parasitic habits throughout the entire group, though possibly some
cases may prove to be an advanced type of predatory habits, differing in this
respect from the other Tabanoidea.

The Tabanidae and Leptidae have amongst their numbers what may be
regarded as the nearest to the primitive venation for this particular superfamily,
but in antennal structures they are considerably advanced and specialized, the
nearest to the primitive type being perhaps that of the genus Pelecorrhynchus.

The Stratiomyiidae, on the other hand, have a very specialized venation as
the radial vein has moved bodily forward towards the costa, which specialization
must not be confused with that of the Nemestrinidae and of the Asiloidea where
only the apical parts of the veins have so moved forward. The most advanced
type in the Stratiomyiidae is to be found in the genus Ophiodesma (Kertesz, 1923,
p. 118 under the name Diapontomyia) where not only has the radial moved forward
but also the median, this vein becoming fused with the radial, the radial-median
crossvein being obliterated in the process. In antennal structures there are many
forms that must be regarded as approaching the ten-segmented primitive type,
and many forms have also preserved the primitive elongate abdomen, where each,
or most, of the segments are as long as, or longer than, wide.

The primitive interradial crossvein seems to have become atrophied in the
Tabanid-Leptid stem, missing in the Stratiomyiid stem and preserved in the
Nemestrinid-Cyrtid stem; it occurs in the genera Cyclopsidea, Trichopsidea and
Exeretoneura, the last of these has also retained a strong trace of the primitive
ten-segmented antennae and moreover it is the only form known to me that
indicates the possibility of its stem having arisen from a type with an elongate
abdomen.

The second superfamily, the Asiloidea, contains insects that do not have a
pulvilliform empodium and apparently have evolved from a somewhat advanced
land-breeding type. This hypothetical type differs from that of the Tabanoidea
by having only seven distinct antennal segments, a larger number has not been
found represented in any existing genus and indeed the occurrence or the traces
of this seven segmented antennae is persistently represented throughout the

382 PHYLOGENY OF DIPTERA BRACHYCERA,

superfamily. The small family Scenopinidae is said to be allied to the Therevidae
owing to the similarity it bears to that family in the larval stages, but th
relationship is very anomalous.

There are two main branches in this superfamily, the Bombyliid stem and
the Asilid stem. The first of these seems to have lost entirely one of the
branches of the median vein but in other respects there is a marked parallel
development to that of the Asilid-stem.

In the Bombyliidae the first radial vein often becomes short, running to the
wing margin much nearer to the base than is usual and this has adlowed the other
branches of the radial to run to the costal margin when they have curved upwards
as is found to be the case in so many genera. The median vein also shows a
tendency to curve upwards as M, runs into R, in certain cases.

In the Asilid stem, the first radial remains normal and when the following
veins are strongly curved upwards they run into this vein and not to the costal

Empididae
\
iw Dolichopodidae

. e
(ey
2
S
Y
; 4
Mydaidae Neuiiesintets
\ ae ' .
NK Bombvliidae ' Cyrtidae
. . | 4
Apioceridae NA . 17
Ne Se A ' (Acanthomeridae)
‘ hod a ~ 6 t ‘
= 1’ Sif, 1 i oh
IN Aue Piles: ; , Stratiomytidae
/ / Ss \ in D I / Ss = a
Asilidae / S I L a
n s -\~
x / \
1 _ Q
Therevidae Scenopinidae ue \ T
‘~~ ~- Tabanid
xy abanidae
ews Leptidae

/
/

Text-fig. 1.

margin. This is a parallel development to that of the Bombyliidae resulting in a
strikingly distinct vein pattern.

All the Mydaidae and Apioceridae, and practically all the Therevidae (the
exceptions may not belong to this family) are provided on the female with genital
spines that are similar in every respect to those of certain groups of the Asilidae,
and which are not found in the Bombyliidae or families other than the Asiloidea.

The third superfamily, Empidoidea, is composed of two families, the Empididae
and Dolichopodidae, both of which contain flies that have larvae associated with
water as is the case with the Tabanoidea, but besides being without the pulvilliform

BY G. H. HARDY. 383

empodium they are also marked by the restriction of the cubital and anal veins,
these being considerably reduced in length.

The diagram (Text-fig. 1) of the relationship so far discussed would conform, I
believe, to something near the average diagram that would be drawn by any
entomologist who had a fairly comprehensive knowledge of the Brachycera, and
to the work so far accomplished in building up the relationships between the
various families. Differences of opinion would centre chiefly about the position of
the Nemestrinid-Cyrtid stem and of the family Scenopinidae, and also concerning
the exact relationships of the Asiloidea families, excluding the Bombyliidae.

There can be little doubt that the spines at the apex of the female abdomen,
spines used for raking the ground when oviposition takes place, are significant
in so far as all those insects that contain them must be considered as derived from
a common stock and that they do not occur outside the Asiloidea suggests that
they originated within the Asiloidea itself probably remote from the deviation
ef the Bombyliid stem which itself has an analogous row of genital bristles that
are used in quite a different manner.

I have stated above that the Asiloidea were derived from a stock having but
seven antennal segments and this is probably correct in the main in so far as
the other segments had largely amalgamated with one or other of these seven.
It is possible that the whole ten may have been indicated by annulations, etc.,
and in building up the structure from the lowest type of antenna in the Asiloidea
I have yet found, that of the Mydaidae, I would unhesitatingly place the first two
segments as 1 and 2, the shaft following as 3 and 4 (these are clearly enough
indicated), the apical flattened sections as 5, 6, 7 and 8, the minute process at the
extreme apex, situated on the edge just within a depression as 9, and finally
the minute spine on this minute process as 10.. These last two are so easily
overlooked, the 5-8 so completely amalgamated that there thus becomes only five
of the original ten segments clearly defined but nevertheless seven are indicated.
The abdomen of the Mydaidae is quite primitive but modified in its type and the
vestige of the interradial is present, but in every other respect the wings have a
very specialized venation. The head is deeply excavated between the eyes very
much as it is on most of the Asilidae. There are plenty of hairs above the oral
opening but they do not form the appearance of the typical moustache of the
Asilidae as they are soft and slender.

The Apioceridae have only four antennal segments clearly defined, the last
being a short and strong spine. The head is slightly excavated between the eyes
as on certain Asilidae, the abdomen is much shortened, and the antennae are
situated too near the oral opening to permit of a moustache being formed there;
the hairs on the very small area so formed are soft and slender. It is evident
that the Apioceridae are very advanced along their line of development, the
venation being specialized and the interradial crossvein is entirely missing.

The Therevidae consists of a number of genera of diverse forms and amongst
their numbers are to be discovered many primitive forms, one specimen that has
a rather long abdomen also has five antennal segments, all easily detected, the
last being a spine, and in addition there is the rudiment (or possibly remnant)
of an Asilid-like moustache; the hairs along the oral margin consisting of a
series of bristles such as is found on certain Asilidae but more slender, comparable
with that of Neocyrtopogon. In no case has there been discovered any form
with the head that is to be compared with the excavated head of the Asilidae
and Mydaidae. With regard to venation, this is fairly simple but invariably lacks

384 PHYLOGENY OF DIPTERA BRACHYCERA,

the interradial crossvein; at most M, runs into M, whilst the cubital and anal
veins may also meet near the apex. The genus Clesthenia is excluded from these
remarks and given special consideration below.

As already pointed out, all the three families, Mydaidae, Apioceridae and
Therevidae are provided with spines at the apex of the female abdomen. This
character occurs in many Asilidae and in view of the similarity of other
characters already referred to, characters that unite these four families, it seems
probable that these genital spines indicate a common origin for all those forms
containing them.

If it were possible to separate the family Asilidae into two or more sections,
one stem of which has arisen from a stock containing these spines, the remainder
from a stock that did not develop them, then it would be reasonable to suppose
that the Mydaidae, Apioceridae and Therevidae had a common origin with one
stem of the Asilidae, which in the present case would be the tribe Saropogonini,
a section of the Asilidae, not the whole stock. It does not seem reasonable to
suppose that the spines could have developed independently and assumed identical
formations as is to be found in the four families.

Saropogonini

Therevidae
Apioceridae
Brachyrrhopalini
Laphrini
Mydaidae
Phellinj
Asilint
Leptogasterini Om matini
Bombyliidae

Text-fig. 2.

The genus Clesthenia was originally placed in the Leptidae by White, but
subsequently removed by me to the Therevidae; I now believe the position here
to be anomalous. It is certainly one of the Asiloidea and shows a marked advance
from the primitive stock. Five antennal segments are discernible, the abdomen is
rather short and conical, the venation lacks the interradial crossvein and M, runs

BY G. H. HARDY. 385

into M,, otherwise the veins are simple. It may be an off-shoot of a more
primitive Asilid type rather than that of the Therevid stem and if so the genus
should prove of exceptional interest. Similarly the American genus Pherocera
Cole, which is said to be without the genital spines, would also need to be placed
elsewhere.

Doubtless these views will be modified by further research, but if they are
adopted for a working hypothesis, it will be necessary to amend the portion of
the genealogical tree relating to the Asiloidea as in Text-fig. 2.

In concluding these remarks I would draw attention to the apex of the female
abdomen which, in the Tabanoidea, is provided with simple cerci in all its three
stems whilst, in the Asiloidea, these cerci have not been found or perhaps are
modified in certain cases such as to provide the lamellae on the female ovipositor
of the Asilini. It is possible other characters of the female genitalia have yet to
be found that would be suggestive in regard to these relationships and so a study
of this aspect should yield some important data.

To the list of works referred to in this paper, there are also added others
that bear upon the subject dealt with and amongst these will be found diagrams
that will serve to illustrate the remarks made above.

Postscript, added 13th September, 1927.

Whilst the above paper was going through the press, a Bombyliid of the genus
Comptosia was found using the bristles at the apex of the abdomen in the
manner identical to that exhibited by the Saropogonini when ovipositing. The
genitalia of this Bombyliid were thereupon studied in considerable detail and the
conclusions drawn were to the effect that the type of ovipositor exhibited By it
approached nearer than any other examined to the hypothetical type from which
the Saropogonini and Therevidae were derived. Moreover this hypothetical type
which can now be defined with some assurance, may have been the origin of all the
widely varying types found in the Asiloidea, for certain features of the Asilini
type are to be traced to it through the genus Neoaratus and the spines of the
ovipositor on the genus Philonicus may yet be shown to have had the same origin
as those on the Saropogonini.

For the immediate purposes of this paper it is necessary to draw attention
to the order in which the relationships of the various families and tribes have
been built up. This was done on the understanding that the primitive type of
ovipositor was represented only by a pair of valves generally greatly reduced, and
the end segments of the abdomen were more or less telescopic as in Mecoptera
(Tillyard, 1926, p. 340). If this be the case, then sorhehow in the Asiloidea at
least, a modification must have arisen resulting in the building up from a simple
to the complex ovipositor that in many cases became permanently exserted. It now
becomes advisable to revise this view and perhaps a better conception of the
phylogeny within the Asiloidea may be gained by completely reversing the order
of the Asilid stem so as to bring the Mydaid-Saropogon branch to the foot where
other characters would ally it, and by placing the Leptogasterini at the top.

References.
Coxe, F. R., 1923.—‘“‘A Revision of the North American Two-winged Flies of the Family
Therevidae,’ Proc. U.S. Nat. Mus., \xii, pp. 1-140.
DaKIN, W. J., and ForpHAM, M. G. C., 1922.—‘‘Some New Asilids from Western
Australia,” Ann. Mag. Nat. Hist. (9) x, pp. 517-530. (Contains two good illustrations
of the genital spines on the female.)

386 PHYLOGENY OF DIPTERA BRACHYCERA.

Harpy, G. H., 1919 and 1921.—‘“‘Australian Rhyphidae and Leptidae,” Proc. Rey. Soc. Tas.,

1919, pp. 117-129; “Australian Bombyliidae and Crytidae,” ibid., 1921, pp. 41-83. i

, 1921, 1924-1927.—“‘A Preliminary Revision of Some Genera Belonging to the
Diptera Brachycera of Australia,’ Proc. LINN. Soc. N.S.W., xlvi, pp. 285-300;
“Australian Nemestrinidae,” ibid., xlix, pp. 447-460; “Australian Mydaidae,” ibid.,
1, pp. 137-144; “A New Classification of Australian Robberflies Belonging to the
Subfamily Dasypogoninae,”’ ibid., li, pp. 305-312; “‘A Reclassification of Australian
Robberflies of the Cerdistus-Neoitamus Complex,” ibid., li, pp. 643-657; ‘Further
Notes on a New Classification of Australian Robberflies,” ibid., lii, pp. 387-398.

IRWIN-SMITH, V., 1920-1923.—‘“Studies in lLife MHistories of Australian Diptera
Brachycera,’’ Part i Stratiomyiidae, Part ii Asilidae, Proc. Linn. Soc. N.S.W., xlv,
pp. 505-530; xlvi, pp. 253-255; pp. 426-432; xlviii, pp. 49-81; xlviii, pp. 368-380.

Kurtesz, K., 1923.—‘“‘Vorarbeiten zu einer Monographic der Nothacanthen,” Ann. Mus.
Nat. Hung., xx, pp. 85-129. (Wing of Ophiodesma under the name Diapontomyia
rufispina n. sp., p. 118.)

MACKERRAS, I. M., 1925.—‘“‘The Nemestrinidae of the Australasian Region,’ Proc. LINN.
Soc. N.S.W., 1, pp. 489-561.

MauiocH, J. R., 1917.—“A Preliminary Classification of Diptera, Exclusive of Pupipara,
Based upon Larval and Pupal Characters, with Keys to Imagines in Certain
Families,’ Bull. Illinois State Lab. Nat. Hist., xii, pp. 161-407.

MELIN, D., 1923.—‘‘Contributions to the Knowledge of the Biology Metamorphosis and
Distribution of Swedish Asilids in Relation to the Whole Family of Asilids,”’
Zool. Bidrag. Uppsala, viii, pp. 1-317.

TILLYARD, R. J., 1926.—‘‘Insects of Australia and’ New Zealand’’—Sydney.

FURTHER NOTES ON A NEW CLASSIFICATION OF AUSTRALIAN
ROBBERFLIES (DIPTERA-ASILIDAE).

By G. H. Harpy.

Walter and Eliza Hall Fellow in Economic Biology,
Queensland University, Brisbane.

[Read 29th June, 1927.]

For loan of, or access to material that aided in the formation of this outline
on a proposed new classification of Australian Asilidae, I would again express my
indebtedness to the Directors and Curators of Museums and to various
entomologists all of whom have been enumerated in an earlier paper (Hardy,
1926, p. 306); for the loan of further material to Mr. Alan Dodd, chief of the
staff engaged upon the biological control of prickly pear, and to his co-workers
Messrs. A. R. Taylor, G. R. Bassingthwaighte and T. A. Cole, who have submitted
for identification genera and species I have not seen represented in other collections.
Those new species from Chinchilla, Queensland, often labelled ‘“’Chilla” in hand-
writing, are liable to lead to errors if they ever find their way into collections
abroad and become described from there; the corruption is too near “Chile” unless
very clearly written.

My knowledge of the world’s genera of Asilids and their structure is mainly
based upon the few papers available to me, to Lundbeck’s Diptera Danica, and to
Melin’s excellent studies on the Swedish species. The last mentioned author
anticipated many of the views I was slowly evolving and so the enlightenment
afforded by him gave me greater confidence than I would otherwise have had in
the classification proposed. To these works I must add an acknowledgment
to the value of the exotic Asilids sent by Professor M. Bezzi, consisting for the
most part of genera related to the Australian forms. I have insufficient knowledge
of the genus Dasypogon itself, therefore I have made no attempt to place it in
the present work; but from what I can gather it will form the typical genus of
a tribe not represented in Australia.

In 1921, I revised the characters of various genera of Australian Asilidae,
and again in two papers of 1926 reconsidered some of these. These three papers
marked a stage in a search for an adequate classification which is further advanced
in the present paper wherein the family is treated as a whole. Three proposed
tribes have already been diagnosed and these are maintained in the present paper,
though a different conception is given to some of them and four more tribes are
added, two of which have already been mentioned.

An improvement will be found in the incorporation of part of the old subfamily
Laphrinae with the Saropogonini, two genera being thus transposed, whilst the
Cerdistus-Neoitamus complex is more adequately isolated though very little altered
in effect, the genus Stilpnogaster being now excluded from that complex.

Vv

388 CLASSIFICATION OF AUSTRALIAN ROBBERFLIES,

Three subfamilies only are recognized, the tribe Laphrini being too intimately
connected with Brachyrrhopala and Phellus to permit of its isolation except
perhaps under quite a different conception to that understood by the old subfamily
Laphrinae. Saropogonini, judging from Australian collections, seems to warrant
subfamily rank, but this is not followed in the present paper. The subfamily
Asilinae is now divisible into two tribes, Ommatiini and Asilini, whilst the
typical genus of the latter, Asilus, is excluded from the Australian fauna.

These tribes are recognizable from the following key:—

ie Amtennaeswith alone nair-likevarisStawat ap Cxsumrerscireieiceeeriiers -)o cs) cicneion sla isiononeasnenetelie 2
Antennae never with an arista, at most with a minute hair-like spine that is usually
situated within a cavity, depression or in a notch. (Subfamily DASYPOGONINAE.) 4

2. Venation very simple, alula of wings much reduced in size; female genitalia simple
and not formed with an ovipositor. (Subfamily LEPTOGASTERINAE.) (One tribe

COW 00 AAD Na trol oped icho- riche mepiotarekeree cache OIGtor cack Ion oto mEEONORD.O.Gc0-070. 0.0. 6: CED ENCE REN LEPTOGASTERINI
Venation with the vein R,,, joining R, before reaching the wing border, also M,
similarly runs to M, (Subfamily ASILINAH.) ...................++.+.+2200- 3

3. Arista of antennae pectinate; female with simple genitalia, not being formed with
BW ChIP goaducoonenneOUoU dda noo Ud GO OOO UKDO KOO oo nO UOebO OND OO 6 OMMATIINI
Arista of antennae simple; female genitalia formed with an ovipositor ...... ASILINI

4. Female abdomen with genital spines at apex. Venation more or less simple, or
R,,, runs into R, or M, runs into M,, but both sets of veins do not coalesce in the

SAIN ISDECIES | ecrepe abs ee reper ete ch ce euromaiioonerets Tekogeha tne Non nsec Ronee eeememewe-tepiey/< etait SAROPOGONINI
Female never with genital spines at apex, but if spine-like hairs occur these are
numerous and do not take the form of strong curved spines as in the Saropogonini

so dco DDdOD OHA OUOND HOOD UOC HOO DOL ODO OODOODUODDOONDD DODO SO UDODABDDDDODDOONONS 5
5. Wings never with M, running into M, ...................-..--.- BRACHYRRHOPALINI
Wings always with M, running into My, ........-...--2-. sees ee eee eee eee ee eee 6
6. Wings never with R,,, running into R, -...-....-..2 2.22 ee eee eee eee eee eee PHELLINI
Wings always with R,,, running into R, ......---.---+. 2 esse e eee ee eee LAPHRINI

The hypothetical genealogical relationship between these tribes is shown in
the following diagram where an attempt has been made to interpret the relations

Brachyrrhopalini Saropogonini

Phellini .

Laphrini AGES

Leptogasterini

Om matiini

BY G. H. HARDY. 389

by the degree of specialization taking into account genital characters of the
female, antennal structure and other outstanding characters. If the isolation of
the Saropogonini is maintained, as illustrated in the diagram, it will be necessary
to erect a subfamily name for the group.

Subfamily LEPTOGASTERINAE.

This subfamily contains but one tribe under which the characters are dealt
with. It conforms nearest to the Asilinae in antennal structure but to the
Brachyrrhopalini in venation, only being more simple in this respect. The
female genitalia are simple as far as I can see, consisting of a vent between the
ninth dorsal and ventral sclerites. Lundbeck refers to a small, generally quite
hidden, ovipositor, and although the ninth segment does not bear the grey
tomentum of the others and hence is black, it does not seem to be specially
shaped into an ovipositor in the Australian forms.

Tribe Leptogasterini (new tribal name).

This tribe contains flies that are very slender and have a much reduced alula
on the wing, thus giving the insect the pedicellated wing structure that is
reminiscent of dragonflies of the suborder Zygoptera. The simple venation with
all the veins running to the wing border without unduly curving, and the absence
of the interradial crossvein, which is but rarely indicated by a stunted vein, the
absence of pulvilli on all legs and the very long claws between which an elongate,
slender empodium occurs, are consistent characters occurring on all species of the
only genus yet recognized from Australia.

Genus LEPTOGASTER Meigen.

Under this genus Melin recorded a resting habit where the flies “swoop down
on a grass-stem with the wings lying parallel in order to rest. In doing so they
usually catch hold of the stem with the tarsi and hang down with the abdomen at
an angle of 45° to the stem (Fig. 14)”. He then quotes Lundbeck: ‘When sitting
the hind legs are generally stretched forward so that all six tarsi are placed near
each other”, adding his own remark, “I have never seen this, on the contrary, the
flies sat as shown in the figure with the back tarsi widely separated from the
anterior pair’. It so happens with the Australian species the temporary resting
position on an upright stem of grass is always as Melin reports, but they are
liable to rest at the extreme apex of a horizontal twig, the body held in a direct
line with the twig which the head almost touches, and are usually sluggish or in
a state of coma when so found. The best time to observe this is in the early
morning and late afternoon, although they may be so found at any hour of the day.
Further it should be noted that the elongate hind legs are quite near the anterior
pair, being placed on the twig just anteriorly to them. It will be readily realized
that the fly in such a position looks so like a part of the twig on which it rests,
that the attitude may be taken as a case of protective mimicry which is heightened
by the elbowing of the legs downwards, these looking like triangular pieces of
cobweb around a notch; an excellent “camouflage”. These observations suggest
that Lundbeck recorded the resting habit when in a state of coma, whilst Melin
observed only the temporary resting periods.

390 CLASSIFICATION OF AUSTRALIAN ROBBERFLIES,

The following species are described from Australia :—

Gonops geniculatus Macquart, 1850.

Leptogaster Leptogaster
pedanus Walker, 1849. vernalis White, 1913.
antipodus Bigot, 1878. - aestiva White, 1913.
australis Ricardo, 1912. fumipennis White, 1913.
bancrofti Ricardo, 1912. occidentalis White, 1918.
dissimilis Ricardo, 1912. autumnalis White, 1916.

Of these eleven names, Walker’s and Bigot’s species have been placed as
synonyms of Macquart’s, although Walker’s name should have taken priority.
In the various collections examined, I can recognize only six distinct species,
three of which are apparently undescribed. The most satisfactory method of
recognizing the various species is by the shape of the male genitalia, but
unfortunately there are few authentic specimens in the Australian collections
and only one type. The usually accepted colour characters, upon which most
species were founded, are quite inadequate for specific determination but may
give a clue to identity, so I sent drawings of the genitalia to Dr. G. A. K. Marshall,
Director, Imperial Bureau of Entomology, asking to have these compared with
specimens in the British Museum collection, expecting in this way to get
satisfactory determinations, and although the drawings were compared, I received
word that they did not correspond to any genitalia in the specimens in the
British Museum. Further notes indicate that the type of L. pedanus Walker has
lost the tip of its abdomen, and there are no males representing L. vernalis White,
fumipennis White and autumnalis White.

Subfamily ASILINAE.

Four papers have already appeared (1920-1926) treating with this subfamily,
mainly dealing with the descriptions of species, but in one (1921), an attempt was
made to deal with generic conceptions and in another (1926) with genera allied
to Cerdistus. In the first preliminary survey the genera were not at all adequately
placed and the relationships were inferiorly dealt with. In the meanwhile it has
become possible to recognize some very marked lines of division making two
groups worthy of at least tribal value.

Tribe Ommatiini (new tribal name).

Ommatius stands alone, having no apparent relationship with any other
genus. The female genital apparatus appears to be more in keeping with the
Leptogasterini, being apparently primitive and in no way comparable with the
highly developed ovipositor type of the other Asilinae.

There is but one genus in Australia which is represented by a large number
of species for which twelve specific names are available. They are all provided
with a pectinate arista which enables both sexes to be easily relegated to this
tribe, and the males have their genitalia of a characteristic shape.

Tribe Asilini (new tribal name).

This tribe contains six genera in Australia that are readily grouped into

three groups of related genera, and diagnosed in the following key:—
Ee, AMLELEAGIA nn CLOSSVeIN BDLESCIIL Lathe Tere hare choice ete) MeN de M ER MEM MORets Gls! o\celtsiiooneltctatte Promachus
Enterradial\ crossvein’ Absent ses ace cts tare oie eM RUAN Cey el ates (ooo Monaite tone eh etceltepelette 2
2. Eighth and ninth ventral sclerite of abdomen clearly defined in the ovipositor ......
pad ah asia) ee V at ate kene"toits ol p hois¥ sxe) apte ants be apauaniel Ou ve te darev et acs rehekel Seema MeRe Ronee Cerdistus, Neocerdistus
Ninth ventral sclerite of the abdomen not discernible in the formation of the
OVIPOSICOR fee. oo OE an ee EE nee Neoaratus, Pararatus, Blepharotes

BY G. H. HARDY. 391

I have recently discovered that a wrong interpretation has been given by me
in previous papers concerning the term “lamellae.” The position where they occur
marks the anus as pointed out by Melin. For the whole structure, when compact,
I used the term “lamella” in the singular where the plural should have been
used, unconsciously regarding the complete organism as a unit, much in the same
way as a segment of the abdomen is singular though composed of two sclerites.
Moreover Melin shows the presence of lamellae in Philonicus, which character was
overlooked by me, the line of division being obliterated on my specimen. Also I
have since found them on Neocerdistus; they were overlooked till a specimen with
the ovipositor buckled was examined in detail, the division between the eighth and
ninth dorsal sclerite being clearly defined on this specimen, as also is the division
between the lamellae when seen dorsally. .

The genera referred to by me (1926) as being allied to the Cerdistus-Neoitamus
complex may be divided now on the following and better characters :—

1. Ovipositor with only one ventral sclerite, the ninth ventral abdominal sclerite not

oe dina Syn Gl] LU Clie evs Shcgi sss: csselte- oe citauelpeuianeMen strets| ta eitemenextaneicek suiausl crieisueel ad Mikey wt aiatsalteriay absevumt totic Mauna irabees sa 2
Ovipositor with two clearly defined ventral sclerites ...............2 00 eevee 4

2. Lamellae with long spines placed dorsally ............. 0... ccc eee ecseeece Philonicus
amelliaemwithoOwtye Su Gly (Sle Siete se bk ie ene cee su suerich a Paes ous gs ER iG octane oon sue repens ee 3

3. The ninth dorsal segment extending above lamellae thus limiting the range of its
movements; i.e. lamellae wedged in .................... Dysmachus, Hutolmus
Lamellae free ...... Heligmoneura, Machimus, Tolmerus, Epitriptus and Stilpnogaster

4, The two upper lamellae fixed, each being produced into a spinelike appendage at the
DDE Ky Ofs sel Sma PG ONS a dA cid ees hy i eA) Seep eee ea ELLOS oe sah Coe ch Vos RSE Oct Neocerdistus

MWAmMeMAC IEC ean GW SiMe nace er eons dei esto p dn een, nome Onicha aan cll SBM Se eS Faye adit ea nib years cris Sat yreuray tah
PRT ene aerate Cerdistus, Neoitamus, Paritamus, Trichoitamus and Rhabdotoitamus

It will be noted that Stilpnogaster is now eliminated from the Cerdistus-
Neoitamus complex and so the name is no longer available for subgeneric purposes.
Machimus is removed from the Dysmachus group, the lamellae being shown as free
in Melin’s illustration; on my specimen it is doubtfully so.

In describing the ovipositor on different species of the Asilinae, Melin wrote
(p. 219): “The dorsal part of the ninth abdominal segment, which. is often more
or less semicircular, is usually drawn out ventral-terminally into a horizontal
lamella which bounds the anus beneath. In consequence of this the terminal
lamellae, even when they are called free, are more or less wedged into the ninth
dorsal segment”. This is quite a justifiable criticism but nevertheless there
appears to be a considerable amount of movement in the lamellae, the lower plate
of which moves in unison with the upper ones and hence must be flexible. This
does not appear to be the case with forms referred to as having the lamellae
“wedged in”.

Genus PromacHus Loew.

This genus is unique amongst Australian robberflies by having the interradial
crossvein forming an apparent continuation of R, This vein runs parallel to the
main stem of R,,, to near where R.,, branches off. In antennal and general
characters it conforms fairly well with other Asilini. The ovipositor is clearly
composed of the eighth and ninth abdominal segments, both ventral and dorsal
sclerites of each being conspicuous. There is sometimes a tendency to incorporate
within the ovipositor the sixth and seventh segments much in the same way as is
accomplished with Neoitamus, but in such cases these segments retain the
tomentose covering. In a specimen before me of P. laciniosus Becker, from Tunis
(Algeria), the eighth segment of the abdomen is covered with hair similar in

392 CLASSIFICATION OF AUSTRALIAN ROBBERFLIES,

quantity and length to that on other segments, but the tomentose covering is
absent, nevertheless it has the general appearance of the anterior segments. The
ninth ventral sclerite is clearly indicated and at first sight appears to be the
only one incorporated in the ovipositor; there are Australian species that approach
this form.

Genus CERDISTUS Loew.

This genus is a complex under which come species having an ovipositor that
is usually very compressed, always with normal free lamellae and the ninth
ventral segment clearly defined. Another genus, as yet only represented in
Australia by two females from Westwood, Queensland, has the appearance of
those belonging to Cerdistus, but is without a discernible ninth ventral sclerite,
the ovipositor is of moderate length, very compressed.

CERDISTUS FULVIPUBESCENS Macquart.

? Glaphyropyga australasiae Schiner, Reise Novara Dipt. 1866, p. 187.

Synonymy.—From the description of Glaphyropyga australasiae, there can be
little doubt that the species should be placed under the complex Cerdistus, but
the German in which the description is written is rather difficult to translate and
even with a fresh description made for me by Mr. F. Rose I have been unable
to satisfy myself regarding Schiner’s meaning in some characters he gives. Miss
Ricardo states that the description reads suspiciously like that of Heligmoneura
lascus Walker a New Zealand species, but should it prove Australian as recorded
then it is likely to be the same as CO. fulvipubescens Macquart. I know of no other
light coloured species that approximates the description so well.

Genus NEOCERDISTUS Hardy.

There is a necessity to amend some characters given for this genus the
reasons for which are incorporated in the discussion above. The ovipositor is
sub-compressed, showing a convex dorsal surface, the ninth dorsal and ventral
sclerites are clearly defined in it, the lamellae are somewhat difficult to detect
in outline, the two dorsal plates are produced, each into a spinelike process that
stand apart from each other; the presence of the ventral plate is problematical.
The first median cell (that formed by the branching of M, and M,) is strongly
constricted subapically. In other respects the genus resembles Cerdistus.

Genus Nroaratus Ricardo.

Including Asilus of Australian authors.

The Australian forms hitherto placed under the genus Asilus (Hardy, 1920,
pp. 250-259) do not belong to it, for the female ovipositor differs fundamentally
from that of the typical Asilus; the habits of oviposition are also quite distinct.
Melin described and illustrated the ovipositor of Asilus crabroniformis which is a
complex of the eighth and ninth dorsal sclerite, the lamellae and the eighth
ventral sclerite. In the Australian forms they vary slightly, but the eighth dorsal
sclerite may be covered with tomentum and therefore similar in colouration to
the rest of the abdomen and in such cases forms a covering from which protrude
the ninth dorsal sclerite and lamellae, and the ventral sclerite which is apparently
of one piece, more depressed than compressed. From this, which occurs on the
typical form there is a tendency to vary towards incorporating the eighth dorsal
sclerite which then becomes black and shining but does not alter in shape to any
extent and therefore never reaches the form found in Asilus.

BY G. H. HARDY. 393

Melin described the supposed mode of oviposition by Asilus crabroniformis;
the ovipositor inserts the eggs between particles of cattle droppings in which
action the short spiny bristles at the apex of the ovipositor are supposed to play
a part, but the Australian forms deposit eggs exposed on a drooping leaf, the
bristles referred to in Asiluws are therefore not required and hence not present;
another distinguishing feature.

I observed the oviposition in a garden at Edgecliff, Sydney, where early one
morning I found N. inglorius Macleay depositing on a leaf of a shrub. Already
one complete orderly row was formed and a second nearly so. There were
several gaps in the second row which were being filled, the lamellae operating as
a feeler for the purpose. The fly later took a step forward and started a third
row this constituting two eggs together, then two further eggs wide apart. Later
when I returned four rows were completed and the fly was gone, but the gaps
in the third row were filled and in all four rows there were only three or four
breaks in the sequence of the eggs, whilst three or four eggs were misplaced.
Early one morning exactly a week later the larvae hatched and fell to the ground.
I was fortunate enough to see this, having been on the look out morning and
evening every day. Miss V. Irwin Smith obtained oviposition in the case of the
typical Neoaratus under artificial conditions in a manner that suggests that that
species has similar egg-laying habits.

Genus PARARATUS Ricardo.

The female ovipositor is well developed and without a ninth ventral sclerite,
but ventrally, at the apex which tapers to a point, there are a pair of flanges
strongly developed. Where such flanges occur on species of other genera they are
small and mainly hidden. The lamellae, composed of three flat plates, are
distinctly defined and not at all compact as on Asilus, Neoaratus and Cerdistus.
Only one species is known, P. macrostylus Loew, recorded from Western Australia,
Queensland and Victoria.

Genus BLEPHAROTES Westwood.

This genus is readily recognized by the broad depressed abdomen containing
conspicuous tufts at the sides. In the ovipositor the ninth dorsal segment is
clearly defined as also the three plates of the lamellae but ventrally only one
segment is defined, this seen laterally, is broadly rounded at the apex.

Under this genus the male genitalia of four species were illustrated in outline
(Hardy, 1921, p. 296), two being described as new. The figures were correctly
given in the text but the names were transposed under the explanation to the
figures. Text-fig. 9 is B. brisbanensis, since found to be a synonym of B. flavus
Ricardo (Hardy, 1923, p. 176); Text-fig. 10 is B. punctatus Hardy; Text-fig. 11
which was left unnamed is the common form found around Sydney to which I now
apply the name B. corarius Wiedemann. Text-fig. 12 is a rare undescribed form
of which I have seen only the male. The readily recognized B. abdominalis is in
nearly every collection. B. vivaxr Hermann is represented by two females only
and there are no further species in any collection examined. The females are
best allied to the males by noting the proportional length of the antennal segments
as these coincide in the sexes, no two species having the same proportions. With
regard to colour both light and dark yellow abdomens occur in three of the
described forms and colour therefore has no significance for identification purposes.

394 CLASSIFICATION OF AUSTRALIAN ROBBERFLIES,

Tribe Phellini.

The tribe Phellini contains but two genera Phellus and Psilozona, both of
which require a more intensive study. The female of a new species from
Chinchilla, Queensland, is in such excellent condition that all the genital parts are
readily recognized and conform in type to no other ovipositor I have seen in the
Dasypogoninae; very distinctly does it show the absence of genital spines such
as found on the Saropogonini. This new form, red and black in colouration is an
undoubted Phellus of which previously I had studied only old specimens;
presumably the spines discovered on them will ultimately prove to be but
modified hairs that have no relation to the spines of Saropogonini.

On Phellus the moustache is very thick and depressed, the hairs all pointing
more or less towards the apex of the proboscis, whilst on Psilozona they conform
to the more usual types, the hairs being upstanding, issuing more or less at right
angles to the face. On P. glaucus and P. piliferus the moustache is overlaid with
a quantity of flattened out scales that arise from the face, expand in width, then
become ribbon-like and finally taper to a point. These specialized hairs are not
found on the specimen from Chinchilla nor yet does the apical prolongation on the
intermediate tibiae occur on that form.

Tribe Laphrini (new tribal name).

This tribe is constituted by that part of the old subfamily Wapainae which
has three antennal segments with or without a hair-like spine but when present
it is placed dorsally as in many Brachyrrhopalini. It is further distinguished by
the veins R, and R,,, amalgamating before reaching the wing border and by the
veins M, and M, doing likewise. Hven when so limited the tribe is complex.

Six genera are placed here, namely: Nusa, Atomasia, Adelodus, Cyanonedys,
Laphria and Maria all of which are recognized in the various collections except

the last. Aphesia has been used for one species which Hermann suggests is an
Adelodus.

Tribe Brachyrrhopalini.

This tribe has already been satisfactorily diagnosed and four genera defined
in key form (Hardy, 1926, p. 307). In none of the species do any of the radial
or the median veins meet each other before reaching the wing margin, but
occasionally M, runs into R,. Most of the species are recognized and suitably
labelled in collections and the status of each species is fairly well assured. The
following is a complete list with synonyms:—

Brachyrrhopala Macquart, 1847 (Cabasa Walker, 1850).
*pulchella Macquart, 1846 (Dasypogon).
venno Walker, 1849 (Dasypogon).
rujfithoraz Walker, 1850 (Cabasa).
rubrithoraz Macquart, 1850 (Dasypogon).
*ruficornis Macquart, 1847.
fenestrata Macquart, 1850 (Codula).
victoriae Roder, 1892.
maculata Roder, 1885.
fulwa Ricardo, 1912.
? quadricincta Bigot, 1878 (Codula).
Codula Macquart, 1850.
*limbipennis Macquart, 1850.
? vespiformis Thomson, 1868.
Chrysopogon Roder, 1881.
*albopunctata Macquart, 1846 (Dasypogon).
spintha Walker, 1849 (Dasypogon).

BY G. H. HARDY. 395

*crabroniformis Roder, 1881.
*mulleri Roder, 1881.
fasciatus Ricardo, 1912.
punctatus Ricardo, 1912.
queenslandi Ricardo, 1912.
*splendissimus Ricardo, 1912.
*rufulus White, 1918.
*nigriscens White, 1913.
pallidipennis White, 1917.
rubripennis White, 1917.

Opseostlengis White, 1913.
*insignis White, 1913.

Note.—Species bearing the names here given with an asterisk may be regarded
as correctly named in collections; doubt may be raised concerning some of the others.

Tribe Saropogonini.

To this tribe now come all forms of the Australian Asilidae that are
provided with strong curved spines in the genital complex of the female, four
or five segments in the antennae, the last of which consists of a minute spine
inserted in a cavity at the apex of the penultimate segment. Two genera,
Clinopogon and Bathypogon are exceptions in so far as the last antennal segment
is of considerable size, being more segment-like and not a hair-like spine. It will
be noted that those species hitherto described fall into two groups on characters
that appear to be suitable for key purposes; those with the tibial spur on the
anterior legs and those without, the latter a complex, the former being again
divisible on the shape of the abdomen though all the forms are apparently
closely related.

For convenience the first group is divided in the key on the presence or
absence of the scutellar bristles as this works well, and it leaves only the genus
Saropogon in an anomalous position. A more natural grouping of the genera
would be accomplished by separating those forms with a tapering abdomen or
allied thereto from those with a clubbed abdomen or related thereto, but these
forms meet towards having the abdomen somewhat cylindrical and it is not
always easy to determine to which category a specimen should be aligned.
Apparently all the described forms with the tapering abdomen or related thereto
have scutellar bristles, only one undescribed form is without them, whilst all
those with clubbed abdomen or related thereto are without bristles except
Saropogon.

The isolation of genera hitherto proposed and which are grouped in the
key given below is offering considerable difficulty as many of them are joined
by related but undescribed species. Even Thereutria and Metalaphria cannot be
isolated satisfactorily for there are two species as yet undescribed that are like
each other in every respect except the veins R, and R,,, are separate in one from
South Australia and run together in the other from Queensland, thus joining
the Thereutria type of wing to the Rachiopogon type.

Key to the genera of the Saropogonini.

1, JANN IOP MnO, Tiida El GurAveul Gobie EE AE Gboogodooucccscaaqcsusonodaesonoaee 2
ANtEHIOMerenlordmwWwAunOUut theN CULVedmESDUE I. selec cece eee ee ae eee 3
2. Always with scutellar bristles; abdomen tapering from base to apex, rarely otherwise
andweshzhthyaclubbed monly ing Sa7;opOg Olle sorick casei eine eiaiaeieieeiorereieiciale cae
areyeneesusrepaneues Questopogon, Rachiopogon, Thereutria, Metalaphria and Saropogon
Never with scutellar bristles; abdomen club-shaped, basal segments always narrower
than some of the more remote ones, rarely quite cylindrical and never tapering
eye eanot one te tek ewes oie ee ta Neosaropogon, Deromyia, Neocyrtopogon and Hrethropogon

Ww

396 CLASSIFICATION OF AUSTRALIAN ROBBERFLIES,

3. Antennae with the apical segment reduced to a hair-like spine ................ 4
AntennaenwithwchenapicalescsimentenocESOMLeqduced prs iat ee eye ene 5

4. Wings without an interradial crossvein .................... Stenopogon, Neodioctria
Whos Vial eho ToaNeOmeChenl GROSKIN Goaooboadblognraonvadhocouosenddc Cryptopogon

5. The veins M, and M, running to the wing border independently ...... Clinopogon
The veins M, and M, meeting considerably before reaching the wing border ..:.....
SC cr OIG OUR CHO 8 ali) ccs aro CHER ESTO Ect ORF Gira Oo fo 6. a Glofel ater ay Bt eear oer coe Bathypogon

Those species that fall into couplet “2” of the key given above may be placed
in their correct genera in many cases by aid of the following temporary key, but
some species which presumably are undescribed and cannot be made to conform
to the genera of this key are at present regarded as belonging to new genera:—

Key to the genera of the Saropogonini that have a tibial spur.

1. Abdomen strongly tapering; scutellum almost invariably provided with bristles .. 2
Abdomen clubformed, at most cylindrical, never tapering, rarely with scutellar
WDTUS LOS) esate ee char ital sjras sah dalsne ony siiah ote re, ec eels hes DeeUeNGer Be 5) ca cin eNreL Tera eI HC SHORES Meee aya. a tarser exe RO aes ar ae 5

Ph VST TR, ENGL IE WOAAOOOES TOXINS) WINS) Vinbales OMNI 5 55g oD Oo Kano DO dORDOD ood CUE 3

Veins R, and R,,, running independently to the wing margin .................. 4
8. Moustache bushy, not confined to the oral margin ....................+. Thereutria
Moustache confined to the oral margin ................+...6- DN va iadstaers, anes Metalaphria

4. Antennae with three segments and a minute spine; moustache confined to the oral

Marein-eiace HPAare ee egos uwiec issue caess Mekene oka tee MSR el ERR Ae tele bel eo Rachiopogon
Antennae with four segments and a minute spine; long soft hairs on face above
TNOUSEA CHIC) Fayence aire ateies aetna doers tha be ner eine ayel do) ee ee Men Re MUS sc ss coves Questopogon

5. Scutellum with bristles; antennae with four segments and a minute spine ..........
BG GF OlOFeR Gai SINCE DLO Na Erich Ch Geo EAC onc eRGUD che cota Mina enas cy 6 Bb -cui G00! D0 ad cue Eeeee Saropogon
Seutellum? without) DriStlesic ks inn essay cee. coeds fekerse | GRRE RRS Lavi cl- ane cathe tee Mecham 6

6. Vein M, meeting M, before Reachineachewwin eal OG Creare teici iene ais Deromyia
Veins M, and M, running independently to the wing border ...................... 7

7. Antennae with three segments and a minute spine. Face prominent, bare, moustache
confinedato) thevoral marsink ere oad sets eee i G.010 Dio Gena eesesG Neocyrtopogon
Antennae with four segments and a minute spine. Face normal .............. 8

8. Moustache confined to the oral margin; face bare .................. Neosaropogon
Face above moustache with slight soft hairs .....................42. Hrethropogon

Genus THEREUTRIA Loew.

Three common species are recognized in collections and in all of them the
moustache differs in colour according to sex; .all have white lateral spots usually
discernible from the second to sixth abdominal segments. 7’. amaricus Walker,
a common Sydney form that extends into Queensland, has the basal half of the
tibiae yellow. TJ. pulchripes White, which is plentiful in the Blue Mountains,
New South Wales, has the basal half of both femora and tibiae brown, which
gives the insect the superficial appearance of a common Laphria. T. pulchra
Schiner has the tubercle and moustache smaller than those of the other two
and the legs are entirely black but white hairs are abundant on the posterior side
of the intermediate and posterior tibiae; it is abundant around Brisbane. The
outstanding name placed here, 7. luctuosa Macquart, may possibly belong to
T. pulchra Schiner.

Genus MeraLaryria Ricardo.

This genus differs from Thereutria only by having the moustache reduced to
a row of bristles along the oral margin. A rare Brisbane species is certainly
congeneric with M. australis Ricardo and may be identical with it. Although
UM. aurifacies White agrees in generic characters it seems to be related more to
Thereutria than here. White’s species is not uncommon around Sydney during
November and December. There is a black species from Chinchilla which has
the abdominal spots of a Thereutria, the moustache of Metalaphria and the wing
venation of Rachiopogon and Questopogon and it is certainly a connecting link.

BY G. H. HARDY. 397

Genus ————————_ ?
Neosaropogon Ricardo in part.

Attention has already been drawn to the fact that Neosaropogon Ricardo,
was a composite genus when established and of those species originally placed
under the name only WN. princeps could be retained there. The other species
have not yet been allied to any genus. D. salinator Walker, N. claripennis Ricardo,
N. nigrinus Ricardo and N. froggatti Dakin and Fordham are all described as
having the tibial spur present and with bristles on the scutellum and hence
must belong to the Rachiopogon-Questopogon group. At least two of these forms
are provided with tapering abdomens that are unusually long, resembling in this
respect the genus Neoaratus, one of the Asilini.

A pair of N. froggatti agrees remarkably well with the description and
conforms to this unnamed genus; the male has quite normally shaped genitalia
with no outstanding features as seems to have been expected by Dakin and
Fordham (1922). WN. claripennis is congeneric, N. nigrinus has not been recognized
and N. salinator is represented by two species so named in Australian collections,
one certainly congeneric, the other doubtfully so.

Genus Saropocon Loew.

Saropogon semirufus Bigot has been recognized and there are two other
species, apparently undescribed, that come here. Saropogon (Dasypogon) suavis
Walker is applied to specimens in Australian collections to a small species of
Neosaropogon, Miss Ricardo having labelled one as doubtfully this. Brachyrrhopala
bella White was removed to this genus tentatively by me; I have not been able
to check its characters since. Saropogon rubescens White, the type of which I
have before me, belongs to the Rachiopogon-Questopogon group; it is smali in
size, but the abdomen tapers strongly. I also suspect that S. dissimulans White
does not belong to this genus, but I have not recognized it.

Typically the genus contains species that have a slightly clubbed abdomen
and scutellar spines and is related to the Neosaropogon group rather closely.

Genus NeEosaropocon Ricardo.

There are now three species described that are recognized as belonging to
this genus, Dasypogon princeps Macquart, D. nitidus Macquart and D. suavis
Walker, but there are quite a number of species in collections that must be
Placed here.

Genus DrromyiaA Philippi.

In Australia this genus appears to be confined to the higher mountains and
the relationship is undoubtedly with Neosaropogon from which it differs by the
veins M, and M, uniting before reaching the wing border.

Genus NrocyrTopocon Ricardo.

I have recently seen the typical form of this genus from Westwood, Queensland,
and find it conforms in every respect to the generic characters previously given
by me, though based upon other species. The whole head is very inflated, the
excavation between the eyes being eliminated; the moustache is confined to the
oral margin and is remarkably reduced in regard to the size of the bristles. It is
further distinguished from Neosaropogon by the entire absence of the fourth
antennal segment. Other species that have been placed by me in this genus are

398 ; CLASSIFICATION OF AUSTRALIAN ROBBERFLIES.

less developed with regard to the head characters, but agree in antennal structure
and therefore cannot be confused with other genera of the Neosaropogon group.
The typical form may be mistaken for a Therevid on structural grounds but it is,
nevertheless, a member of the Saropogonini.

References.

DAKIN and ForDHAM, 1922.—‘“‘Some New Asilids from Western Australia,’ Ann. Mag.
Nat. Hist. (9) x, pp. 517-530.

Harpy, G. H., 1920.—‘‘Descriptions of Australian Flies of the Family Asilidae,’ Proc.
LINN. Soc. N.S.W., xlv, pp. 185-202; and “On the Male Genitalia of Robberflies
Belonging to the Subfamily Asilinae,”’ ibid., pp. 250-259.

, 1921.—“‘A Preliminary Revision of Some Genera Belonging to the Diptera
Brachycera of Australia,’? Proc. LINN. Soc. N.S.W., xlvi, pp. 285-300.

, 1923.—‘‘Notes on the Distribution of Some West Australian Asilids,’’ An.
Mag. Nat. Hist. (9) xii, pp. 174-176.

, 1926.—“‘A New Classification of Australian Robberflies Belonging to the
Subfamily Dasypogoninae,” Proc. LINN. Soc. N.S.W., li, pp. 305-312; and “A Re-
classification of Australian Robberflies of the Cerdistus-Neoitamus Complex,” ibid.,
pp. 643-657.

IRWIN-SMITH, V., 1923.—‘‘Notes on the Egg Laying, Bggs and Young Larvae of
Neoaratus hercules Wied.”, Proc. LINN. Soc. N.S.W., xlviii, pp. 375-380.

LUNDBECK, 1908.—Diptera Danica, Part 2, pp. 1-87.

MELIN, 1923.—‘Contributions to the Knowledge of the Biology, Metamorphosis and
Distribution of the Swedish Asilids in relation to the Whole Family of the Asilids,”’
Zool. Bidrag. Uppsala, viii, pp. 1-317.

TILLYARD, 1926.—‘‘The Insects of Australia and New Zealand” (‘The system of naming
the veins adopted in the present paper is the amended Comstock-Needham notation
taken from this work, pp. 337-338).

Proc. Linx, Soc N.B.W., 1927,
1 Prare

©
A RECONNAISSANCE TOPOGRAPHIC SURVEY
— of.

——-THE KOSCIUSKO PLATEAU — ~~

lie su

ete eecearnay Aes Eas eet oe sae s

Bayh Saks “ne cals

a Se * ay Sas ie, CaO ee ;
2 RS ae yi yet % Re

Proc. Linn. Soc. N.S.W., 1927. PLATE XI.

MouNT KOSCIUSKO.

1. Slope of a shallow valley at about 5,100 feet. H. coriacea consociation at rear.
Poa consociation on the slopes at sides.
2. Shallow valley at about 5,500 feet, showing destruction of H. coriacea.

Proc. Linn. Soc. N.S.W., 1927. PLATE XII.

Mount IXoscrusko.
1. View at 5,200 feet, showing H. coriacea in the background and Callistemon in the
foreground.
. Restio associes at 5,500 feet.
. Shallow valley at 5,600 feet with EH. coriacea on the ridge and Poa in the foreground.

to bo

Proc. Linn. Soc. N.S.W., 1927. PLATE XIII.

Mount Kosciusko.
J. Granite boulder at 6,000 feet with B. Gunniana flattened against it.
2. V. Derwentia in a dead E. coriacea forest at 5,500 feet.

Proc. Linn. Soc. N.S.W., 1927. PLATE XIv.

MouNT KOSCIUSKO.
1. Regenerating plant of C. Sieberi.
2. Snowy River, at 4,000 feet with H. Gunnii in the background and Leptospermum
lining the river bank.

aoa
ee

iv
i
af

: ;
.
:
ve
: rs
i
: aityes Pr
.
prs * ’
Pepe at 6
i ven: A
WE LR cs
, "Wl'tp ‘phe rou
ry '
Hi AM,
: i
e j
i

Proc. Linn. Soc. N.S.W., 1927. PLATE XV.

ue

COME Pitas :

Mount KOSCIUSKO.

1. Ep. paludosa in flower in middle distance, E. coriacea in background, 5,100 feet.
2. Ep. paludosa in flower, with Poa-Hypolaena passing into the Poa society.

Proc. Linn. Soc. N.S.W., 1927. PEATE XVI.

Mount KOSCIUSKO.

Alpine region at 6,700 feet.
Ep. paludosa at 5,100 feet.
E. Gunnii at 4,000 feet.

Shallow valley at 5,800 feet.

Tr
ber

ie
Pyaiiler ae
i tts
fps al Ny on
iy. eee
iar ea

Proc. Linn. Soc. N.S.W., 1927. PLATE XVII.

Mount KOSCIUSKO.

1. Poa-Hypolaena ecotone developing into Epacris associes.
2. Ridge in slate area at summit of plateau occupied by Ep. petrophila.

Proc. Linn. Soc. N.S.W., 1927. PEATE, Xvi

MouNT KOSCIUSKO.
E. coriacea at 5,200 feet showing regeneration after fire.
Poa-Celmisia association at 7,000 feet.

Ale
2
3. Alpine region seen from summit, 7,328 feet.

Fate
Eta

*

Proc. Linn. Soc. N.S.W., 1927. PLATE XIX,

MouNT KOSCIUSKO.

ra

Podocarpus alpina in the shelter of a boulder at 7,000 feet.
Scene at 6,000 feet.
8. Celmisia in flower at 7,000 feet, with dense societies of Huphrasia Brownit.

RS

SEG dpe aN Shy

Fane pal easte

-*

Proc. Linn. Soc. N.S.W., 1927. ; IPN) SS,

Australasian s:ecies of Disciseda.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXI.

Australasian species cf Disciseda and Abstoma. Spores of Disciseda.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXII.

Mycenastrum coriwm. Spores of Disciseda, Abstoma and Mycenastrum.

tarive

fi
by

any

a

Tt ie

Proc. Linn. Soc. N.S.W., 1927. PLATE XXIII.

Sections of leaves of Hakea.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXIV.

Sections of leaves of Hakea.

XXV.

ATE

124e,

65 UAT

Soc. N.S.W

N

Proc. Lin

we

cern eet

oe

Y

DIYDET JO S9AVIT JO SUOTIDEg

.
a ee |
RVI 7

‘Mh
Pike

PLATE XXVI.

Proc. Linn. Soc. N.S.W., 1927.

Australian Hesperiidae.

VA,

' ie = Looe ‘ ot i det a Ss a te
; : Wha ke ene
: ¢ :
4 ;

ag oe

NOTES ON AUSTRALIAN DIPTERA. No. xiii.

By J. R. MALiocH.
(Communicated by Dr. I. M. Mackerras.)

(Fifteen Text-figures.) Bi
[Read 28th September, 1927. ]

In this paper I present descriptions of a number of species sent to me by
Mr. A. L. Tonnoir, mostly from Tasmania, the type specimens of which are to
be deposited in the Cawthron Institute, New Zealand. The paratypes, which I am
permitted to keep, will be sent by me for disposition in some suitable Australian
museum. In the paper I also present some keys for the recognition of genera
and species in the Acalyptratae, the types of the new species in most cases being
from material sent to me by the late Dr. E. W. Ferguson, and these will be returned
to Australia later.

Family Sapromyzidae.

Though there are I believe some genera of this family still to be discovered
in Australia I am presenting now a generic synopsis which will I hope permit
students of the group to locate material they may become possessed of, provided
it belongs to an included genus. About half of the genera are confined to Australia
and most of the species are.

Key to Genera.

1. Hind tibia with two strong curved black apical ventral spines, the longest one
over twice as long as apical diameter of the tibia; costa without short stiff
black spines, fine haired from base to apex of fourth vein; hairs on cheeks,
thorax, femora, and tibiae, long and bristly; anterior orbital bristles incurved
LL ie HOR RMR cheba eliss 3)(oas ener spel sus ueveues tie rouehauste eceneceseys gene 1. Amphicyphus de Meijere.

Hind tibia with, or without, one short, almost straight, apical ventral bristle, which
is not longer than the apical diameter of the tibia; part of costal vein with short

closelysplacedmstifiablacks bristles iiyorassmcte crcieticide ole cic eiciele cc cle sieceleler el avaiane 2
2. The short stiff black bristles on costal vein not extending to apex of third vein,
gradually discontinued beyond apex of second ................-..-eseeeee 3

The short stiff black bristles on costal vein extending to apex of third vein where
they cease abruptly, giving the costa the appearance of being slightly thickened

COME AR DO LIN iia feist said Poise ole afebeyseval ela d sSevenaevatabarene?’ Opole s Slelepe-w Mlarerele osaveyevelocetens 15
Second wing vein unusually close to costal vein, the cell (marginal) between these
veins therefore not one-third as wide just beyond apex of first vein as the one
(Submarsinal) behind sSecondaveimise crysis) -iel-lcuclebecienelsictencieyic sicrenelieiclnelatisiieieiens 4
Second wing vein at the normal distance from costal vein, the cell between these
veins just beyond apex of first vein usually as wide as the one behind second
NIGER so: 52e!g BLOC LOIOIGKa Ch SRA DI ERONR Ic EOC RETESET RRA? IPE PPI cers GU et COMA 5 BUS CL OD ete) Van re Beh Ba 5

4. Anterior orbital bristles incurved; basal segment of antennae quite evident in profile,
at least as long as second, and with some fine hairs at apex below; inner cross-

vein of wing at or slightly before middle of wing .... 2. Steganopsis de Meijere.
Anterior orbital bristles curved backward; basal segment of antennae not exposed

in profile; inner cross-vein at about one-third from apex of wing ..............

058.017 10 OID1O OO. DCO) GASH che NC SHDICRCRORGNT ICIS EI MORO ICEL OIG ioe roioie oisue horek 3. Depressa Malloch.

(JX)

400 NOTES ON AUSTRALIAN DIPTERA, Xiii,

De eosthumeralebristlemotupresentaonmthoralxainnne hierar oe een eee 6
Posthumeral bristle generally strong, always present ................00e0ee0ee 1

6. Anterior orbital bristles incurved; cheek bristled ...... 4. Trigonometopus Meigen.
Anterior orbital bristles curved backward; cheek haired ...... 5. Paranomina Hendel.

ei, ANGE joWbO Ci Choke! LOMAS WMA! Gooccaccosoadcduonsusdobscouodnbeon
Anterior pair of orbital bristles curved backward, or indistinguishable .......... 10

8. Thorax with four pairs of strong dorsocentral bristles, the anterior pair in front of
SUCUTOM olicieiciewsisuepsaeteneee emetic ted aes ata: ont Ga MUS frame std tee! Cal mn ie 9
Thorax with two or three pairs of dorsocentral bristles, the anterior pair behind
FSD ADU one mieten ty Seamer sis yf DIST OR HE CRARORS CLAY 6) OFS oan eee toy aioe 6. Incurviseta Malloch.

9. Face glossy, quite prominently convex on upper half, depressed below; arista white
except jal Dalse: ein ie ess nec opeebeneeer tel = 7. Poecilohetaerella Tonnoir and Malloch.

Face not, or but slightly shining, almost flat; arista entirely dark ...............
BCG ERG cle otGs oe CRONOT NS HE OREO OREO REP Sa EE REE eee nen i aestce ans La 8. Poecilohetaerus Hendel.

10. Frons with but one well developed pair of bristles, the anterior pair lacking or
aAlMOSt ein Gistin BUSH a lS yes nye stcrces ce ees ic cerned ee Coo ee a Ta rat nS OEIC HSE oy RSI a cae 11

Frons with two distinct pairs of orbital bristles ....-............0000cceeeeees 12

11. Arista lanceolate on basal half, and with very short dense black hairs which give
it a more thickened appearance; face quite prominently convex on upper half,

with a median impression which is very distinct when seen in profile ........

Be UE «CM PA WEE Babe Aa td, 2) fob Sa et eMac Va Oe ea Ab Se Oe a a 9. Ceratolauxania Henrdel.

Arista not exceptionally thickened on basal half, and with normal pubescence; face
not prominently convex on upper half, if convex then regularly so and not
impressed at middle when seen in profile ............ 10. Paralauxania Hendel.

12. Eye much, head slightly, longer than high, face receding below ................
EP AES RE Pee OR EH BR cde wut Re ee ee 11. Trigonometopsis Malloch.

Eye and head not conspicuously longer than high, the former usually almost

TAG D0 V6 We eerere ema Men Nar eau ar eter ee ee Re AE Re en No giinls ohare dle tiatt G.0.0 6 0 0 13

13. Face except the parafacials glossy black, and prominently, subconically, convex
( GB =p-< rey dtr) Ua eg Ree esta elo seacuta ts 6 ichtaaunc ts ce Sachin tua ate ae RSPR Ch as 12. Melanina, n. gen.

Face not glossy black, and very little or not at all convex .................... 14

14. Frons in profile projecting beyond anterior margin of eye about half as far as
length of eye; basal antennal segment longer than second; ocellar bristles

Vel rebates ol Sire oti quarbechSeasalard esdlaidrd ora cerb/a orckovelolo o ora dd on omar 13. Rhagadolyra Hendel.

Frons in profile projecting in front of eye much less than half the length of eye;

basal antennal segment almost invariably much shorter than second ..........
Be SNE Beet CRS OO DURE O os ORTH Oe STORER EOS Oa LG atch oor asar LOIS OADNO ae 14. Sapromyza Fallen.

15. Mesopleura with a strong downwardly directed bristle near middle of disc, in
addition to the one on hind margin .... 15. Trypaneoides Tonnoir and Malloch.
Mesopleura with only fine hairs on disc, no strong bristle present ............ 16

16. Face slightly or not at all convex; frons little if any longer than wide; lower part
TE OCOie MORMAI! oooodcoooUPecoO Os Dd GOO SOOO 16. Homoneura van der Wulp.

Face with a prominent subconical central prominence; frons about three times as
long as wide; lower part of occiput with a tumid spot close to eye on which

there is at least one strong bristle .................. 17. Australina Malloch.

1. Genus AmMPpHICYPHUS de Meijere.

I have recorded the occurrence of reticulatus (Dol.), the only species of this
genus, in Australia (These Procerpines, 1926, p. 550). There are a few slight
differences in the markings of the single Australian specimen I have seen and one
IT had from Java, but not sufficient to cause a doubt as to their identity.

2. Genus STEGANOPSIS de Meijere.
So far I have seen but one species of this genus from Australia, which I

identify as melanogaster (Thomson).
I have a key to the known species of this genus now ready to send to press.

3. Genus Drepressa Malloch.
I erected this genus originally for the reception of one species, atrata Malloch.
Mr. Tonnoir has sent me two additional species which I describe below, I present
in the following key diagnostic characters of the three species.

BY J. R. MALLOCH. 401

Key to the Species.

1. Face glossy black, but slightly yellowish on upper margin, the black colour extending
over anterior two-thirds of cheeks; fore coxae and bases of fore femora yellow,

mid and hind femora fuscous; costal margin of wing narrowly hyaline from base

to the hyaline preapical fascia, the costal vein pale yellow on same section;
longest hairs on arista three or four times as long as its basal diameter ......

D200 S:-8 G:0-0 070 10'O' 6 01G-O1OEH, ORD ACRES CLOT LOGI AROL NO cc Oa ORAL eE Sac chee Cio Hite ene albicosta, n. sp.

Face largely or entirely fulvous yellow; legs not coloured as above; costal margin
not hyaline from base to preapical fascia; longest hairs on arista about twice as

loneRaseitstbasaliidiameters Crvsrarcuctssueterctoyel carchone ets: snallone et cee ane tenant seas AER ee 2
2. Thorax almost entirely black; mid and hind femora and bases of their tibiae black;
no conspicuous pale streaks along middle of cells of wings .... atrata Malloch.

Thorax yellow on lateral margins of mesonotum and entire pleura; mid and hind
legs yellow; a conspicuous hyaline streak along middle of most of the cells of
wings and one along costal vein from apex of first vein to the preapical hyaline
LAS CLAN er iee cy oredsn si meee ais) se ticauees sida. cu.) due, avensh Aesuaaten es yehste ape) elleveuewensas striatipennis, n. sp.

DEPRESSA ALBICOSTA, 0. SD.

Male.—Head glossy black, centre of frons reddish in front, upper part of face
slightly yellowish, occiput with a broad yellow streak from middle to lower margin
of cheek behind each eye; antennae fuscous, third segment fulvous yellow; palpi
black. Thorax black, yellowish round prothoracic spiracle and on pleural sutures,
the pleura more shining than mesonotum, the latter and the scutellum quite
distinctly shagreened or alutaceous. Abdomen shining black, not shagreened. Legs
stramineous, mid and hind coxae, apices of fore femora and practically all of mid
and hind pairs, fore tibiae except bases, and all of fore tarsi, black, bases of mid
and hind tibiae slightly browned. Wings blackish, with a slightly diagonal fascia
beyond middle just apicad of inner cross-vein and over lower extremity of the
outer one much as in atrata, but there is a quite noticeable hyaline streak extending
from the fascia along costal vein to humeral vein which is not evident in atrata,
tips of wings hyaline, costal vein yellow from base to fascia. Knobs of halteres
black.

Ocellar bristles microscopic; longest hairs on arista over half as long as
width of third antennal segment. Thorax with three pairs of postsutural dorso-
central bristles, the anterior pair weakest, a moderately long pair of prescutellar
acrostichals, and four series of intradorsocentral hairs. Fore femur without an
anteroventral comb; fore tarsus longer than fore tibia, and as thick as it.
Venation as in atrata.

Length, 3-5 mm.

Type, Killara, N.S.W., 29th Oct., 1921 (A. Tonnoir).

DEPRESSA ATRATA Malloch.

This species is about the same size as albicosta, but has the head almost
entirely fulvous yellow, only the ocellar spot being black, the antennae and palpi
are entirely fulvous, the legs are more extensively blackened, there is no evident
white or hyaline margin to the costa, and the dorsal surfaces of thorax and
scutellum are not noticeably shagreened.

Originally described from Sydney, N.S.W. I have before me a specimen from
National Park, Tasmania, 6th Dec., 1922 (A. L. Tonnoir).

DEPRESSA STRIATIPENNIS, N. sp.
Female—Head fulvous yellow, glossy, ocellar region fuscous, along each
margin of each orbital plate there is a brown or fuscous suffusion; parafacials
white dusted as in the other two species; antennae fulvous yellow; labrum and a

402 NOTES ON AUSTRALIAN DIPTERA, Xiii,

large mark on each cheek entirely glossy black; palpi yellow at bases, black at
apices. Thorax dull fulvous yellow, mesonotum and scutellum brownish or
fuscous. Abdomen fuscous. Legs coloured as pleura, fore tarsi blackened except
at bases. Wings fuscous, with the same oblique hyaline fascia as in the other two
species, but there are narrow hyaline streaks along the middle of most of the
cells, and one from apex of first veinsto near apex of second along hind margin
of costal vein. Halteres brown. It must be noted that the type specimen is
teneral and in all probability mature individuals will have the thoracic, abdominal,
and leg colours darker in part and more contrasted with the pale parts than is
the case here.

Structurally similar to atrata, the hairs on arista not nearly half as long as
width of third antennal segment, but the dorsal surfaces of thorax and scutellum
are evidently shagreened.

Length, 4-5 mm.

Type, Mt. Wilson, N.S.W., 19th Nov., 1921 (A. L. Tonnoir).

4. Genus TRIGONOMETOPUS Meigen.

Some time ago I described a species which I placed in this genus. It does not
agree with the genotype, but there are so many species in this immediate vicinity
in the family that are distinguished by characters of chaetotaxy and structure of
the head that I hesitate to add another genus to the list for the reception of one
species. Consequently this species, fuscifrons Malloch, may be allowed to remain
in this genus pending an elucidation of the family in this region.

5. Genus ParanomMina Hendel.

I have not seen this genus, which was erected for the reception of one species,
unicolor Hendel, described from Cape York.

6. Genus IncuRVISETA Malloch.

This genus was originally erected for the reception of maculifrons (Macquart),
but there are many other species which, though not agreeing absolutely with the
genotype in certain characters that might bé considered as of at least subgeneric
value, appear to find their closest affinities here. I present below a synopsis of
the species now available to me, and indicate distinctions in structure, both in the
Synopsis and descriptions, which may justify further division of the group.

' Key to the Species.

1. Frons yellow, with a pair of deep velvety black marks on the interfrontalia .... 2
Frons either with only the ocellar spot black, the remainder yellow, or almost entirely

10) F: Ke) nt en CER OES en GEER Reh a CRE A ama CREO EET 15 SCO 6 ANU CRE REMC Is HKG CHORES Glo ib hea 4

2. Face prominently convex and glossy, central part readily visible in profile; entire
antennae, aristae, and palpi deep black ............... maculifrons (Macquart)

Face flat, or but slightly convex, not, or very slightly, visible in profile, the surface
almost dull; antennae and palpi not entirely black ....................0- 3

3. Apices of all femora and tibiae blackened, those of mid and hind femora very
TIATT OWA, SOM Neve so vals vege onole vayetoMerous thie’ cue ayaa lone e Kee woe rdsu eReR: Can oM Tc kev eususkev affinis, Nn. sp.
Apices of none of the femora and tibiae blackened .................. wilmoti, n. sp.

4. Frons largely black, yellow only on the anterior portion of the triangle, orbital
stripes and triangle glossy on their entire length and extending to anterior margin
of frons, the narrow intervening and lateral areas velvety black; face prominently
convex, readily seen from the side, and glossy yellow, the parafacials densely
white dusted; wings luteous, quite noticeably infuscated at bases; thorax,
abdomen, legs, except the bases of mid and hind tarsi, and the knobs of halteres,
SHINS RDIACKA tetas re ce te ete crate ale oe tee ete a ene eaten OTN STG Neco toes aie flaviceps, n. sp.

10.

iL

BY J. R. MALLOCH. 403

Frons yellow, only the ocellar region blackened, triangle and orbits not conspicuously
differentiated on their entire length, and not extending to anterior margin of
frons; face rarely convex; species not coloured as above, the legs more
extensively yellow and the halteres usually pale coloured ..........-+++.+-. 55

Basal two segments of antennae yellow, third entirely deep black; thorax testaceous
yellow, with two linear black vittae along the lines of dorsocentrals which extend
from anterior margin to well beyond middle; intradorsocentral hairs in four

OTH CCM Pa EI Neyo e leo Ce re eueneicuer oe Teu sash che rs ccevapeutichg ones isle, lene tener sis vittigera, Nn. sp.
Antennae not conspicuously bicoloured, third segment never deep black; thorax not
coloured as above, if yellow then without linear black dorsal vittae ...... 6

Intradorsocentral hairs in but two series and rather strongly developed; thorax and
abdomen fuscous, with a pronounced green tinge, both greyish dusted, the
thorax more densely so and with the humeral angles yellow; apices of palpi

darkened; face quite noticeably convex and glossy in centre .... biseriata, n. sp.
Intradorsocentral hairs in at least four distinct series which are more or less regular,
SOMATA Vey oe Ayavel Tae) SegnoscooosccscanpuonosanodcoboDoDdaMbGOCE 7

Thorax with three pairs of strong dorsocentral bristles, the anterior pair close to
suture, and one pair of strong prescutellar acrostichals which are conspicuously
differentiated from the very short and fine intradorsocentral hairs which do not
closely approach them; halteres fuscous ...............---- tasmaniensis, n. sp.

Thorax with two distinct pairs of dorsocentral bristles and sometimes a weak pair
anterior to them which is not close to suture, the prescutellar pair of acrostichals
sometimes not much longer than the hairs immediately in front of them, the
latter usually quite prominent and approaching close to the prescutellars .... 8

Intradorsocentral hairs very fine and short, in six series; thorax fulvous yellow,
shining, with a poorly defined greenish dorsocentral vitta which suffuses disc of
scutellum also; palpi blackish at apices; anterior sternopleural well developed
Pee retire Maa EeR Stop aetat eRe e te GE) oN cliaa cate otrat debra eves ianeiichremmielsuiel io ejienienieuletlsliersetTatn fulvoviridis, n. sp.

Intradorsocentral hairs in four series, usually quite long, and always rather
conspicuous; thorax not coloured as above; palpi usually yellow .......... 9

Thorax and legs entirely fulvous yellow, the former usually with one pair of the
acrostichals in front of the prescutellar pair quite long and strong; anterior
sternopleural bristle of moderate length; face slightly concave in profile; abdomen
usually yellowish at base, metallic blue-green beyond ........ flavipalpis, n. sp.

Thorax always largely black; legs usually partly blackened ................-.. 10

Face quite conspicuously glossy and convex in centre; palpi yellowish testaceous
ET eR erie cries chal oho semaetanial lcci seetemeraiier cre tehe evclcl she leseuers ter etertetepstietten stealer viridana, Nn. sp.

Face flat or slightly concave, very slightly or not at all shining ......-......... isl

Palpi brownish or fuscous; head not broader than thorax; abdomen brilliant metallic
(GH RANSN ) GCococnasoogenododecooed ooo SoD O DOU GOMoMUD OO HOUR viridula, n. sp.

Palpi yellow; head distinctly broader than thorax; abdomen glossy black, without
GUTNAGE TAACAININE TOMO, SSS gochicticacuc cd donb dooodaoboouO pon latifrons, nN. sp.

Text-fig. 1. Incurviseta affinis, apex of abdomen of male from side.
Text-fig. 2. Melanina plebeia, head from side.
Text-fig. 3. Sapromyza ocellaris, wing.

INCURVISETA MACULIFRONS (Macquart).
I have seen specimens which I refer to this species from various localities in

Tasmania. The characters cited in the above synopsis ought to serve to distinguish
it from the two next described species, which are similarly coloured and have the

404 NOTES ON AUSTRALIAN DIPTERA, Xiii,

distinctive frontal markings. The protuberant face of maculifrons, however,
ought readily to identify the species. The hypopygium is similar to that of
afinis, but the fore femora are sometimes entirely pale, and at best only browned,
the mid and hind femora are not blackened at apices, the apices of all tibiae are
black, as are the apices of mid and hind tarsi, and usually all of the fore tarsi.

Length, 3-4-5 mm.

Localities—Magnet, Burnie, Mt. Wellington, Mt. Field, and Cradle Valley,
Tasmania (A. M. Lea and A. L. Tonnoir).

INCURVISETA AFFINIS, n. Sp. (Text-fig. 1.)

This is the species I previously identified and recorded as maculifrons.

Both species are testaceous yellow, with the ocellar spot, two large, elongate,
and sometimes subtriangular, marks on frons, apices of tibiae and of tarsi, black.

In addition to the characters listed in the key affinis has the abdominal tergites
each with a large black or fuscous mark on each side, including the apical one
(Text-fig. 1), but sometimes, as in the Seaford specimen, the abdomen appears to
be entirely blackened. The wings in both species are yellowish hyaline, and the
aristae in both are almost bare.

The close similarity of those two species and my evident misidentification of
the one described by Macquart is but another argument against identifications of
imperfectly described species except on the basis of type examination.

Length, 3-3-5 mm.

Type, Sydney, N.S.W., 30th Nov., 1924. Paratypes, Mooni, near Cotts, 14th
Febr., 1925; Seaford, Victoria (G. F. Hill); and Burnie, Tasmania, 31st Jan., 1923
(A. L. Tonnoir).

INCURVISETA WILMOTI, 0D. SDP.

This species is very similar to affinis, but the two black frontal marks are
not so extensive, the black paired spots on abdomen are quite small and are
confined to tergites 3 to 6 inclusive, and the legs are entirely yellow. Both these
species have the four series of intradorsocentral hairs stronger and more regular
than is the case in maculifrons, the outer two series in the latter being less
complete and finer than the inner two. Only females of this species are available
to me now.

Length, 3 mm,

Type and one paratype, Wilmot, Tasmania, 8th Jan., 1923 (A. L. Tonnoir).

INCURVISETA LATIFRONS, N. Sp.

Female.—Head testaceous yellow, the frons more brownish and entirely shining,
face white dusted, not shining; antennae.and palpi testaceous yellow, arista fuscous.
Thorax shining black, fulvous yellow on propleura, anterior margin, and on two
vittae which extend along lines of dorsocentrals the whole length of mesonotum.
Abdomen glossy black, with very faint indications of a bluish cast. Legs black,
tarsi testaceous. Wings distinctly yellow, most noticeably on costa, the veins
yellow. Calyptrae greyish, with black margins and fringes. Knobs of halteres
black.

Head wider than thorax; frons about two-fifths of the head width and about as
wide at vertex as long, slightly narrowed anteriorly; orbits not differentiated,
anterior orbitals farther from eye than posterior pair; ocellars widely divergent,
about as long as the postverticals; space between bases of antennae almost as great

BY J. R. MALLOCH. 405

as distance of either antenna from eye; third antennal segment not over 1:5 as long
as wide; arista with very short pubescence; face almost flat; cheek with fine
marginal hairs which extend some distance upward on sides of face. Thorax with
two pairs of postsutural dorsocentral bristles, four series of intradorsocentral
hairs, a pair of prescutellar acrostichals, two sternopleurals, and the propleural
quite strong. Fore femur without an anteroventral comb; all tibiae with a distinct
preapical dorsal bristle. Inner cross-vein at: about two-fifths from apex of discal
cell; ultimate section of fourth vein about 2-5 as long as penultimate.

Length, 5 mm.

Type, Wahroonga, Sydney, N.S.W., 31st Oct., 1926.

This species has the head wider, and the eyes more divergent below when
seen from in front, than any other species of the genus now known to me.

INCURVISETA FLAVICEPS, Nl. SD.

Female.—Shining black, anterior extremity of central part of frons, the face,
and cheeks, fulvous yellow, antennae a little darker; parafacials white dusted;
bases of tarsi of mid and hind legs testaceous yellow. Wings brownish hyaline,
slightly infuscated at bases. Calyptrae and halteres fuscous.

Frons a little longer than wide, triangle and orbits sharply differentiated, all
three extending to anterior margin of frons, the triangle narrow and about
parallel-sided on its apical third or more; postvertical bristles shorter than the
ocellars; anterior orbitals a little farther from eye than posterior pair; face
quite prominently convex in centre, glossy; cheek about as high as length of
antenna. Thorax with the two pairs of postsutural dorsocentrals rather short,
the series continued forward as rather strong hairs, the intradorsocentral hairs
in two series and rather long, posteriorly but little shorter than the prescutellar
acrostichals; anterior sternopleural present; scutellum slightly flattened above.
Abdomen stout, very weakly bristled. Legs as in latifrons. Inner cross-vein a
little beyond middle of discal cell.

Length, 3 mm.

Type, Cradle Valley, Tasmania, 20th Jan., 1923 (A. L. Tonnoir).

The very prominently convex face and high cheeks might justify one in
erecting a subgenus for the reception of this species but there is nothing to be
gained by adopting this course without a fuller knowledge of the group.

INCURVISETA VITTIGERA, Nl. SDP.

Female.—Head testaceous yellow, ocellar region with a narrow fuscous
triangle; third antennal segment and aristae black; palpi yellow. Thorax
testaceous yellow, a little darker on dorsum, and with two narrow blackish vittae
along the lines of dorsocentrals from anterior margin to the anterior pair of
postsutural dorsocentrals. Abdomen discoloured in type specimen, more brownish
yellow than thorax. Legs testaceous, apices of fore and hind femora and tibiae,
and of all tarsi, blackened. Wings hyaline. Halteres yellow.

Frons a little longer than wide, orbits a little shining, the anterior pair of
bristles farther from eye than posterior pair, all bristles of moderate length;
face slightly convex; cheek fully as high as width of third antennal segment.
Thorax with two pairs of postsutural dorsocentrals and a rather strong setula
representing a third pair, four series of short intradorsocentral hairs, one pair of
prescutellar acrostichals, and only one distinct sternopleural. No preapical
dorsal bristle distinguishable on hind tibia. Inner cross-vein a little beyond middle

406 NOTES ON AUSTRALIAN DIPTERA, Xiii,

of discal cell; ultimate section of fourth vein about twice as long as penultimate
section.

Length, 3-5 mm.

Type, Mt. Wellington, Tasmania, 9th Nov., 1922 (A. L. Tonnoir).

Structurally very similar to maculifrons, but the face is not so distinctly
convex, and if the absence of the preapical dorsal bristle from the hind tibia is a
normal character, the species is readily distinguished.

INCURVISETA BISERIATA, N. SD.

Male and female.—Head testaceous yellow, ocellar spot, upper half of occiput
except on sides, aristae, and apices of palpi, fuscous; frontal orbits very slightly
shining; face very distinctly shining in centre, the parafacials dull. Thorax black,
with a greenish tinge, humeral and propleural regions testaceous yellow, the entire
surface with rather dense greyish dust. Abdomen coloured as thorax, but more
noticeably green tinged, and less densely grey dusted. Legs testaceous yellow,
blackened on most of fore femora, apices of hind femora and of fore and hind
tibiae, less distinctly on mid tibiae, and on apices of tarsi, the fore pair almost all
black. Wings hyaline. Halteres yellow.

Frons very slightly convex, all bristles distinct, the anterior orbitals closer
together than the posterior pair; face quite prominently convex; cheek about as
high as width of third antennal segment; arista with very short pubescence.
Thorax with three pairs of postsutural dorsocentrals, the anterior pair short, the
intradorsocentral hairs in two series, setulose and quite long, and the anterior
sternopleural not developed; scutellum convex. Preapical dorsal bristle on fore
and hind tibiae long and fine, the one on fore tibia fully as long as the basal
segment of fore tarsus. Inner cross-vein very slightly beyond middle of discal
cell and a little before apex of first vein; ultimate section of fourth vein twice as
long as penultimate.

Length, 3-3-5 mm.

Type, male, and allotype, on same mount, and originally mounted in copula,
Eaglehawk Neck, Tasmania, 15th Nov., 1922; paratypes, male and female,
Launceston, Tasmania, 29th Oct., 1922 (A. L. Tonnoir).

INCURVISETA TASMANIENSIS, N. sp.

Female.—Head coloured as in biseriata, but the palpi almost entirely black.
Thorax entirely black, shining, with a slight aeneous tinge, the dorsum slightly
greyish dusted. Abdomen glossy black, with a more or less pronounced blue-green
tinge. Legs black, knee-joints and bases of tarsi testaceous. Wings brownish
hyaline. Knobs of halteres fuscous.

Head as in biseriata. but the face is slightly concave in profile, and the
cheek is not as high as width of third antennal segment. Thorax with three
pairs of outstanding postsutural dorsocentrals, the anterior one close to suture,
and one pair of prescutellar acrostichals, the other hairs very short and fine, the
intradorsocentrals in six series, but not continued to prescutellar acrostichals;
both sternopleurals quite prominent; scutellum convex. Legs as in Dbiseriata, but
the preapical dorsal bristle on mid and hind tibiae shorter. Inner cross-vein
usually a little beyond level of apex of first vein.

Length, 3-3-5 mm.

Type and one paratype, Cradle Valley, Tasmania, 17th Jan., 1923; two
paratypes, same locality, 10th Jan., 1923 (A. L. Tonnoir).

BY J. R. MALLOCH. 407

INCURVISETA FULVOVIRIDIS, Nl. Sp.

Female.—Head almost fulvous testaceous, frons slightly and almost evenly
shining, the orbital stripes hardly distinguished, ocellar spot, aristae, and apices
of the palpi, fuscous; third antennal segment slightly darkened above. Thorax
fulvous yellow, distinctly shining, and slightly whitish dusted, the dorsum with a
faint greenish central suffusion or vitta which extends over disc of scutellum.
Abdomen bright metallic blue-green. Legs fulvous yellow, slightly darkened on
apices of fore tibiae and fore tarsi. Wings yellowish hyaline. MHalteres yellow,
knobs brownish.

Frons a little wider than long, anterior orbitals quite small and fine, and
separated by less than twice the distance either is from eye; face quite noticeably
concave in profile; cheek about as high as width of third antennal segment.
Thorax with only two pairs of postsutural dorsocentrals, the anterior pair much
shorter than the posterior, one pair of prescutellar acrostichals, six series of short
fine intradorsocentral hairs, and both the sternopleurals strong; scutellum convex.
All tibiae with a preapical dorsal bristle, the hind pair with the short hairs near
apex on posteroventral side denser than usual, and erect. Inner cross-vein slightly
before middle of discal cell and apex of first vein; last section of fourth vein not
over 1:5 as long as penultimate section.

Length, 3 mm.

Type, Cradle Valley, Tasmania, 16th Jan., 1923 (A. L. Tonnoir).

INCURVISETA FLAVIPALPIS, 0. SD.

Male.—Similar to the last preceding species in colour, the head, thorax, and
legs being almost fulvous yellow, and the abdomen metallic blue-green. The
antennae and palpi are, however, entirely pale, and there is no greenish central
suffusion on the thorax.

Structurally the two species are very similar, but the frons is about as long
as wide and more shining, the thorax has a short anterior third pair of post-
sutural dorsocentrals, all the hairs longer, the intradorsocentral series four in
number and usually a pair of those in front of the prescutellar acrostichals quite
long, while the anterior sternopleural is shorter. Inner cross-vein at or very close
to middle of discal cell; ultimate section of fourth vein about twice as long as
penultimate section.

Length, 3-3-5 mm.

Type, Geeveston, Tasmania, 7th Dec., 1922; paratypes; two, Eaglehawk Neck,
Tasmania, 22nd to 27th Nov., 1922; one, Burnie, Tasmania, 5th Oct., 1922
(A. L. Tonnoir).

INCURVISETA VIRIDANA, DT. Sp.

Male.—Head fulvous testaceous, parafacials paler, dull; frontal orbits shining,
face glossy; arista and ocellar spot fuscous. Thorax fuscous, with a greenish tinge,
humeral angles, propleural and postalar regions, yellowish, the whole slightly grey
dusted. Abdomen metallic blue-green. Legs fulvous testaceous, rather obscurely
darkened on most of fore femora, apices of fore tibiae and tarsi, and apices of hind
femora and tibiae. Wings hyaline. Halteres yellow.

Frons subquadrate; orbits a little differentiated, the anterior bristles quite
long, separated by a distance about four times as great as that of either from eye;
face quite prominently convex; cheek as high as width of third antennal segment.
Thorax with two pairs of postsutural dorsocentrals and a weaker pair in front of

408 NOTES ON AUSTRALIAN DIPTERA, Xiii,

these; intradorsocentral hairs quite long, in four series, the outer two series
not regular; anterior sternopleural bristle short but distinct. Outer cross-vein a
little beyond middle of discal cell and distinctly before apex of first vein; ultimate
section of fourth vein twice as long as penultimate section.

Length, 3:5 mm.

Type and paratype, Mt. Wellington, Tasmania, 25th Nov., 1922 (A. L. Tonnoir).

INCURVISETA VIRIDULA, 0. sp.

Very similar to viridana, but the palpi are blackened at apices, the thorax
has only two distinguishable pairs of postsutural dorsocentrals, and the four series
of intradorsocentral hairs are quite regular and the hairs are short and quite even.
The face is a little concave instead of prominently convex.

Length, 3-3-5 mm.

Type, Cradle Valley, Tasmania, 24th Jan., 1923; paratype, Fern Tree, 10th
Nov., 1922 (A. L. Tonnoir).

7. Genus PoOECILOHETAERELLA Tonnoir and Malloch.

This genus was described for the reception of some New Zealand species and
no representative of it has been seen from either Tasmania or Australia.

8. Genus POECILOHETAERUS Hendel.

This genus was originally erected for the reception of an Australian species,
decora Schiner, which was renamed schineri by Hendel. The species was
originally recorded as from New Zealand but it does not occur there. There is a
second species referred here, punctifacies Tonnoir and Malloch, which occurs in
New Zealand.

9, Genus CERATOLAUXANIA Hendel.

This recently erected genus contained originally only tetanocerina Hendel,
but I have now before me a second species, from Tasmania.

The genus is very similar to Paralauxania as represented by the Australian
forms, but is distinguished from it by the lanceolate arista, which is densely short
haired.

I have not seen the genotype, but it is evidently distinct from the present
species.

CERATOLAUXANIA TASMANIENSIS, Nn. Sp.

Male.—Head fulvous yellow, glossy, much shrunken in the type specimen which
is immature, but the following colour markings are evident: ocellar spot and
orbital stripes darkened, a black spot between each antenna and eye, with white
dusting below it; third antennal segment and arista fuscous. Thorax shining
black, fulvous anteriorly. Abdomen shining black. Legs black, apices of tibiae,
and the tarsi, testaceous yellow, fore tarsi apparently darkened from before tip of
first to tip of fourth segment. Wings hyaline, bases narrowly black. MHalteres
black.

Ocellar bristles short; anterior orbitals lacking; third antennal segment not
as much tapered as in genotype, almost imperceptibly narrowed apically, about
three times as long as its basal width; vertical bristles long. Thorax with two
pairs of postsutural dorsocentrals; scutellum convex. Inner cross-vein at middle

BY J. R. MALLOCH. 409

of discal cell and below apex of first vein; penultimate section of fourth vein
about three-fifths as long as ultimate section.

Length, 4 mm.

Type, Mt. Wellington, Tasmania (A. M. Lea).

In tetanocerina Hendel the third antennal segment is browned at apex only,
the thorax is blackened on dorsum only posteriorly, the legs are black-brown,
with the tips of the tibiae and tarsi reddish yellow. This species was described
from Victoria.

10. Genus ParRALAUXANIA Hendel.

I have placed some Australian species in this genus because they agree with
the genotype in having but one pair of orbital bristles. I have recently examined
the genotype and though it is a duller coloured species than those from Australia
it has no very striking characters by means of which it can be distinguished from
them apart from colour and the presence of dusting on the face. I propose to
retain the generic name for the two species already included in this genus from
Australia and for some others now dealt with, but at some future time
when the relationships of the species of the family are better known it may be

that some other disposition may have to be made of the Australian species or some
part of them.

Below I present a key to the species as I have aligned them.

Key to the Species.

1. Thorax fulvous yellow, with three black dorsal vittae, the median one continued to
near apex of scutellum, the laterals not reaching posterior margin of meso-
notum, and with only two series of intradorsocentral hairs, one on each lateral
margin of the central vitta, the notopleural suture, and a large triangle on sterno-
pleura blackish; abdomen metallic blue-green; legs yellow, fore femora, tibiae,
and tarsi deep black, the fore tarsi thickened .............. atrimana, Nn. sp.

Thorax largely black, or if fulvous yellow then not marked with black vittae, and
always with at least four series of intradorsocentral hairs; fore tarsi not notice-

Ak? Wuoclwneel cocgootoasasovd0ob cobs FOO ODOdC Odo ODDO COOMA ODDO ON DOO OCOD 2

2. Thorax fulvous yellow, slightly darkened on middle posteriorly, «on dise of scutellum,
and along notopleural suture, and with three pairs of strong postsutural dorso-
central bristles, the anterior pair close to suture and without strong setulae in
front of it; mid and hind femora dirty yellow .......... fiavipennis, Nn. sp.

Thorax with the greater part of its surface blackened, and with two or three pairs

of postsutural dorsocentral bristles, when three pairs are present the anterior
pair is much shorter than the others and quite far from suture, with some
loro GIES ihn Woe Ot We Goooweoobovoodcvoo Doc UdU Dd oGaDoa0D ODO cop UDO DOC 3

Mid and hind femora yellow; frons longer than wide, shining fulvous yellow, with

narrow fuscous lines along both sides of the orbital plates, across anterior margin

Co

of frons, and down its centre ...........-. cece eee eee eet eee ees scripta, n. sp.
All femora deep black; frons of variable width, but never marked with fuscous lines
BIS) ihn ADGTO SOGGIEE EUBEcascanoodsaccuduoadcodcono DDN oOD DO DOD OO DU SUOOMOOOE 4

4. Frons distinctly longer than wide, shining, uniformly fulvous yellow, with only a black
ocellar spot; no dark mark at upper extremity of each parafacial close to
antennal bases, but a quite distinct one at lower extremity of each against the
eye; ocellar bristles extremely small and fine; palpi entirely bright fulvous yellow
CPM MRR Me SCR Hol eRe caustic uel eeitehie eke feltsisau(elje} y's) ahah oseicaltekehemopehe/isiie'e)\e\'0/(s fulviceps Malloch.

Frons blackened on more than the ocellar spot; parafacials with a dark mark at
upper extremity close to bases of antennae ....---.- +e see eres eee eters 5

5. Frons widest at anterior margin where it is about 1:5 as wide as its length at middle;
ocellar bristles widely divergent and not half as long as the postvertical pair;
parafacial with an upper and a lower black mark; wings quite conspicuously
blackened across anal angles; fore metatarsus whitish, blackened at apex ......
Re ey eT eepey an PES A cai J adanct tis lay e vakemet cat elahcuale lode eteitstepetcke ¢ felons ie! sie elevata Fabricius.

410 NOTES ON AUSTRALIAN DIPTERA, Xiii,

Frons subquadrate; ocellar bristles quite large, as long as the postverticals; parafacial
with a dark mark at upper extremity close to base of antenna, but without one
at lower extremity; wings blackish at bases, but without a noticeable black mark

ACTOSS' ANAl) TAN SLES aoe shoe a) ste oh ees are sh eh os edealiasiel nl stisyareneuegeaisiene wears onebeuE eel eneucHees Sonne 6

6. Fore tarsi entirely black; palpi fuscous at apices .........:....... nigrimana, n. sp.
Fore tarsi black, basal three-fourths or more of the basal segment testaceous; palpi
ALAUON NS Ce Ooo Geo Cbd Do OGG On mad COM OO De AOC Oo mou pom d OD flavipalpis, n. sp.

PARALAUXANIA ATRIMANA, N. SD.

Male and female.—Head fulvous yellow; frons glossy, ocellar spot black; face
glossy, parafacials rather densely white dusted; inner mouth margin with a dark
transverse line; antennae and palpi fulvous yellow; arista fuscous. Thorax
shining fulvous or orange-yellow, with three black vittae, the central one carried
over the scutellum almost to apex, the lateral pair not extending to hind margin
of mesonotum, a blackish vitta along the notopleural suture, and a large triangular
fuscous mark on the sternopleura. Abdomen brilliant metallic blue-green, the
sternites yellowish. Legs concolorous with thorax, fore pair except the coxae
deep black. Wings yellowish hyaline, faintly darkened at hases behind. MHalteres
dark brown at apices.

Frons a little longer than wide; ocellar bristles about half as long as the
postvertical pair; anterior orbitals not distinguishable; face slightly convex, and
receding a little below; antennae large, third segment tapered slightly to apex
and over three times as long as its basal width; arista with extremely short
pubescence; cheek about as high as width of third antennal segment. Thorax with
two distinct pairs of postsutural dorsocentral bristles and a series of setulae in
front of them, the posterior one of the series rather strong, one pair of strong
prescutellar acrostichals, two series of intradorsocentral hairs, the anterior sterno-
pleural very weak, and the propleural bristle minute; scutellum slightly elongated.
Legs normal, but the fore tarsi much thicker than in any other species of the
genus known to me, the basal segment fully as thick as apex of tibia. Inner
cross-vein at middle of discal cell.

Length, 4-4-5 mm.

Type, male, Wahroonga, Sydney, N.S.W., 31st Oct., 1926; male paratype, Hagle-
hawk Neck, Tasmania, 23rd Nov., 1922; allotype and two paratype females, Cradle
Valley, Tasmania, 11th, 13th, 16th Jan., 1923; one female paratype, Mt. Wellington,
Tasmania, 29th Nov., 1922 (A. L. Tonnoir).

PARALAUXANIA FLAVIPENNIS, Nn. Sp.

Female.—Head fulvous yellow; frons shining, ocellar spot black, subtriangular;
parafacials white dusted, slightly browned opposite bases of antennae; face shining;
antennae and arista black, third segment of former a little reddish at base; palpi
black, yellow at extreme bases. Thorax shining fulvous yellow, slightly darkened
in centre of hind margin and on disc of scutellum, and with a dark vitta on noto-
pleural suture. Abdomen glossy bluish black. Legs fulvous yellow, apices of fore
femora, most of fore tibiae, and all of fore tarsi, apices of hind tibiae, and of mid
and hind tarsi, obscurely browned. Wings deep honey yellow, brown at extreme
bases. Knobs of halteres brown.

Frons a little wider than long; ocellar bristles much longer than the post-
verticals and posterior orbitals; anterior orbitals represented by a pair of
microscopic fine hairs; face rather sharply convex vertically; receding below in
profile; cheek higher than width of third antennal segment; antennae not as long
as in atrimana, the third segment tapered apically and about 2:5 as long as its

BY J. R. MALLOCH. 411

basal width; arista subnude. Thorax with three pairs of strong postsutural dorso-
central bristles, the anterior pair close to suture, one pair of strong prescutellar
acrostichals, the surface hairs quite short, the intradorsocentral series six in
number, and the anterior sternopleural and propleural bristles long; scutellum
semicircular. Legs normal, fore tarsi not noticeably thicker than the other pairs.
Inner cross-vein close to middle of discal cell; outer cross-vein and ultimate section
of fourth vein slightly sinuate.

Length, 4-5 mm.

Type, Cradle Valley, Tasmania, 17th Jan., 1923 (A. L. Tonnoir).

A stronger development of the anterior pair of orbital bristles might incline
one to place this species in the genus Sapromyza.

PARALAUXANIA SCRIPTA, N. SD.

Male.—Head fulvous yellow; frons shining, with a central fuscous vitta which
is widest behind, a fuscous line around the orbital plates except posteriorly, and
one across anterior margin above antennae which connects with the dark mark
on upper extremity of each parafacial; face shining; parafacial white dusted, with
a dark mark at upper and another at lower extremities; antennae and palpi fulvous
yellow; aristae fuscous. Thorax blackish, with a green lustre, becoming fulvous
anteriorly. Abdomen metallic blue-green. Legs dusky fulvous yellow, the fore
pair more obscured, but without sharply defined markings. Wings yellowish
hyaline, more yellow anteriorly. Halteres fuscous.

Frons a little longer than wide; ocellar bristles widely divergent and about as
long as postvertical pair; anterior orbitals represented by weak microscopic hairs;
face convex above in middle, depressed near mouth; cheek about as high as width
of third antennal segment; antennae smaller than atrimana and flavipennis, third
segment but little tapered apically and about twice as long as its width at base;
arista subnude. Thorax with three pairs of postsutural dorsocentrals, the anterior
pair weak and short, well behind suture, four series of rather long and unevenly

spaced intradorsocentral hairs, the anterior sternopleural and propleural bristles
rather long.

Length, 4 mm.
Type, Cradle Valley, Tasmania, 23rd Jan., 1923 (A. L. Tonnoir).

PARALAUXANIA FULVICEPS Malloch.

Since describing this species I have seen additional specimens from Wahroonga,
Sydney, N.S.W., 24th Oct., 1926.

PARALAUXANIA ELEVATA Fabricius.

Mr. Tonnoir has sent me a specimen of this species from Adventure Bay,
Tasmania, 29th Dec., 1922. Originally the species was not definitely recorded
from any locality, but Hendel gives it from Nova Cambria and New South Wales.
I have seen only Tasmanian specimens so far.

PARALAUXANIA NIGRIMANA, Nl. Sp.

Female.—Head fulvous yellow; frons shining, rather extensively blackened
round ocelli and on anterior extremities of orbital plates; parafacials white dusted,
with a brown mark at upper extremity of each; face shining; antennae largely
infuscated, third segment palest below basally; arista fuscous; palpi black at
apices. Thorax shining black, yellow on humeral angles and propleura, and with

412 NOTES ON AUSTRALIAN DIPTERA, Xiii,

slight greyish dust. Abdomen glossy black. Legs black, mid and hind tarsi
testaceous at bases. Wings yellow, blackened at bases, the dark shade hardly
showing on anal angle. MHalteres black.

Frons slightly depressed near front, and about as long as wide, entirely glossy;
ocellar bristles as long as postverticals; anterior orbitals represented by micro-
scopic hairs; antennae as in scripta; face slightly convex; cheek as high as width
of third antennal segment. Thorax as in scripta, but the four series of intra-
dorsocentral hairs are shorter and quite regular. Inner cross-vein at middle of
discal cell.

Length, 5 mm.

Type, Cradle Valley, Tasmania, 12th Jan., 1923 (A. L. Tonnoir).

PARALAUXANIA FLAVIPALPIS, N. Sp.

Male and female.—Similar to elevata and nigrimana in general colouration,
but the palpi are entirely yellow, and the wings are not conspicuously blackened
at bases, the anal angle being almost without a dark suffusion.

Like nigrimana, a more slender species than elevata, with narrower frons,
shorter third antennal segment, and the characters cited in the foregoing key to
species. The anterior pair of orbitals are entirely lacking in this species.

Length, 4 mm.

Type, male, Wahroonga, Sydney, N.S.W., 31st Oct., 1926; allotype, same
locality, 24th Oct., 1926; female paratype in poor condition, Jenolan Caves, Nov.,
1910 (J. B. Cleland).

11. Genus TRIGONOMETOPSIS Malloch.
I have seen but one species of this genus, the genotype, binotata Thomson,
which appears to be not uncommon in New South Wales.

12. Genus MELANINA, novum.

This genus is similar to Sapromyza Fallen in the thoracic and wing characters,
but the head is of entirely different structure, the frons being more or less
depressed in front, entirely shining, with the orbital stripes a little more highly
polished, the anterior pair of orbital bristles is smaller than the posterior pair,
and are backwardly curved, the other bristles are all present, the face is highly
polished and very distinctly convex (Text-fig. 2), arista pubescent; thorax without
any presutural dorsocentrals. Genotype, Melanina plebeia, n. sp.

Key to the Species.

1. Inner cross-vein of wing at about one-third from apex of discal cell; apical two
segments and a part of third of mid and hind tarsi fuscous, the rest pale
yellowish testaceous; frons quite broadly yellowish in front, the anterior pair
of orbital bristles very small and weak; only one pair of prescutellar acrostichals
developed; scutellum with microscopic pile on entire surface .... major, n. sp.

Inner cross-vein of wing at or very close to middle of discal cell; only the apical
segment of mid and hind tarsi fuscous, the basal four segments pale yellowish
testaceous; frons usually almost entirely black or very narrowly yellowish in
fronts whe Ancerior. Pair oOL OLrpitalDLiStles GIStiN Comin icant ielenioneone 2

2. Only one irregular series of bristles besides the acrostichal two series, i.e., the intra-

dorsocentral hairs in four series; scutellum with microscopic pile on entire surface,
not appearing nMighly polished ‘on! diseh 27% . 3) sere einele creel es ies quadriseriata, n. sp.

Intradorsocenutral hairs Ini SIRVSELICS iE eRe hel a cuceruemeten aie hone) ehaireld ities yoleytted taal atetroReVen 3

Scutellum with microscopic pile on margin only, highly polished on disc; colour

deep glossy black; thorax with one or two of the pairs of intradorsocentral
hairs developed bristle-like in front of the prescutellar acrostichals ..........
Ae dS ER tile, SRE Bas es ts MELD NPRM Ach bte tnd charntece cores ie ane MD LEOETAS | Nae!

w

BY J. R. MALLOCH. 413

Seutellum with a shagreened or dusted appearance and not highly polished on disc,
under a very high power lens appearing as microscopically pilose; colour metallic
blue-black or greenish black, the dorsum of thorax noticeably greenish posteriorly,
and the face distinctly purplish or violaceous when seen from above; all the
HACK OG UIE! VENTE Ga Goocadedodcodpaobedoooocugwuobooc aenescens, n. sp.

All the species are very similar in colour and structure, being distinguished

essentially by the characters cited in the foregoing key. The general colour is
glossy black, sometimes with a metallic blue or green tinge, the legs are black,
with bases of the mid and hind tarsi testaceous yellow, the wings are yellowish
hyaline, sometimes with the bases more or less distinctly infuscated to apices of
the basal cells, the knobs of the halteres are black, antennae entirely fulvous
yellow, and the aristae and palpi fuscous. In the brief descriptions below only the
differentiating characters are noted.

MELANINA MAJOR, DN. SD.

Male.—Deep glossy black, the frons bright orange yellow in front, becoming
brownish above middle, but the interfrontalia distinct from the glossy black
orbital stripes and the ocellar spot on all parts when seen from in front. The
intradorsocentral hairs are short and in six series.

Length, 3-25 mm.

Type, Wahroonga, Sydney, N.S.W., 24th Oct., 1926.

MELANINA QUADRISERIATA, 1. Sp.

Male.—A deep glossy black species, which is rather more slender than the
others in the genus and has the frons almost entirely black. The other characters
may be gleaned from the Key.

Length, 2-5 mm.

Type, Wahroonga, Sydney, N.S.W., 17th Oct., 1926.

MELANINA PLEBEIA, 0. Sp. (Text-fig. 2.)

Male and female—Similar in colour and general habitus to quadriseriata,
but the intradorsocentral hairs are in six series, and the frons is distinctly yellow
on anterior margin. The hypopygium of this and of the next species are similar.

Length, 2-5-3 mm.

Type and six paratypes, Wahroonga, Sydney, N.S.W., 17th to 24th Oct., 1926.

MELANINA AENESCENS, 0. SD.

Male and female.—Distinguished from plebeia by the characters cited in the
key. The frons is rather more distinctly yellowish on anterior margin and the
third antennal segment is less than 1:5 as long as wide, while in plebeia it is rather
more than 1:5.

Length, 2-5-3 mm.

Type and eight paratypes, Cronulla, N.S.W., December, 1925 (H. Petersen).

13. Genus RHAGADOLYRA Hendel.
This monobasic genus is unknown to me. The genotype is handlirschi Hendel.

14. Genus SaproMyza Fallen.

I have before me several undescribed species of this genus, but consider it is
not an opportune time to present a new key to the species as I have but recently
published one in These ProcrEpINGs (1926, p. 34). Under each species described
in this paper I discuss the characters by means of which the species is separated

414 NOTES ON AUSTRALIAN DIPTERA, Xiii,

from the one to which it runs in the key, and believe that with these notes in hand
students may be able to identify reliably such species as are already described.
There are yet many species of the genus to be discovered and every lot received
by me has in it species unknown to me, so that any key presented now will of
necessity prove quite inadequate for the identification of a host of species occurring
in Australia, and especially those from the west and north, which are very
sparingly represented in the material available to me at this time.

SAPROMYZA PLUMISETA, N. SD.

Male.—Frons orange yellow centrally, orbits, vertex, and triangle, fuscous,
whitish dusted; face and cheeks testaceous yellow, with greyish dust, parafacial
suture slightly darkened, a dark spot between each antenna and eye; occiput
fuscous, grey dusted; antennae fulvous yellow; aristae and the hairs black; palpi
fuscous. Thorax fuscous, more yellowish on sides anteriorly, dorsum with dense
grey dust, two practically complete submedian vittae between acrostichals and
dorsocentrals, two narrow interrupted vittae laterad of the dorsocentrals, and two
partial sublateral vittae behind the suture; disc of scutellum suifused with brown.
Abdomen blackish, distinctly shining. faintly grey dusted. Legs testaceous, at
least the fore femora darkened. Wings greyish hyaline, with a broad costal
fuscous cloud which begins at apex of auxiliary vein and extends to tip of wing,
becoming obsolete posteriorly in the submarginal and first posterior cells on disc
of wing, and not connecting with the more conspicuous fuscous clouds over the
inner and outer cross-veins nor the fainter one over bases of second section of
third and fourth veins, but suffusing entire apex of wing to behind fourth vein.

Frons about 1:25 as long as wide; orbits well differentiated, anterior orbital
bristles closer together than posterior pair; all frontal bristles long; third
antennal segment about 1:5 as long as its basal width; arista plumose; face
sharply convex. Thorax with 1+3 dorsocentrals and 1+4 acrostichals; scutellum
flat above. Fore femur with an anteroventral comb; all tibiae with preapical
dorsal bristle; mid tibia with two apical ventral bristles. Inner cross-vein at two-
fifths from apex of discal cell, and noticeably beyond apex of first vein; ultimate
section of fourth vein about twice as long as penultimate section.

Length, 3 mm.

Type, Woy Woy, November, 1921 (A. L. Tonnoir).

SAPROMYZA PETERSENIT, 0. Sp.

Female.—Head testaceous yellow; interfrontalia brown, becoming orange-red at
anterior margin; orbits, and sides of triangle behind, densely whitish grey dusted,
the former as wide as interfrontalia in front and extending to eyes; a deep brown
mark between each antenna and eye, and another on occiput behind each eye
which does not descend below level of eye; upper part of cheek silvery white
dusted; face with a slight dark mark in each antennal fovea; antennae testaceous,
third segment and aristae missing in type; palpi fuscous. Thorax fuscous, yellowish
on humeri, densely pale grey dusted, mesonotum with two narrow dark brown
vittae between the acrostichals and dorsocentrals, which fuse at posterior margin
and continue as one almost to apex of the scutellum, and two much broader sub-
lateral vittae of same colour; pleura with a pale-dusted vitta extending back-
ward from prothoracic spiracle. Abdomen fuscous, slightly shining, densely grey
dusted, each tergite except the first visible one with a dark brown irregular basal
fascia. Legs testaceous, fore femora, and a subbasal and apical ring on each tibia,

BY J. R. MALLOCH. 415

fuscous. Wings hyaline, with a blackish cloud from apex of auxiliary vein to
tips which extends inward over second vein at base, then over third vein from
beyond the cloud over inner cross-vein, connects with a fuscous mark on middle
of apical section of fourth vein, and extends round apex of wing to fourth vein,
but does not connect with the clouds on the cross-veins; a dark cloud over the
veins above base of discal cell, and fainter marks in second postericr and anal
cells. Halteres yellow. a

Frons about 1:5 as long as wide; ocellar bristles almost as long as frons;
anterior orbital bristles very little farther from eyes than posterior pair; frons
protuberant in front in profile; face carinate above, slightly receding below; cheek
a little higher than length of frons in front of eye; antennae broken off in type.
Thorax with bristles as in the preceding species. Fore femur with an antero-
ventral comb; all tibiae with a short preapical dorsal bristle; mid tibia with
one apical ventral bristle. Inner cross-vein at about two-fifths from apex of discal
cell.

Length, 3 mm.

Type, Cronulla, N.S.W., December, 1925 (H. Petersen).

This species, which is named in honour of the collector, is placed with some
doubt in Sapromyza as the third antennal segment is lacking in the type. The
specimen was received from Dr. C. F. Baker, of the Philippine College of Agricul-
ture, in a miscellaneous lot of insects and fragments of flowers, etc., collected by
sweeping and shipped dry in a mass in a tin box. A number of other species
described by me were received in the same manner.

This and S. plumiseta belong to a group in which there are four pairs of
strong dorsocentral thoracic bristles and distinct wing markings present. I have
described only one such Australian species, suffusa (These PROCEEDINGS, 1926,
p. 37), and from it these two may be distinguished by the presence of an antero-
ventral comb on the fore femora, and the strong biseriate acrostichal setulae.
Possibly both species have the aristae plumose but it is impossible to say as to
peterseni; suffusa has the arista pubescent.

SAPROMYZA HIEROGLYPHICA, N. SD.

Female.——Head testaceous, densely greyish yellow dusted; frons with a
blackish brown spot at base of each bristle and a streak on each side from base of
inner vertical to ocellar bristle which runs forward alongside of the narrow
grey triangle to anterior margin, and then sends a branch diagonally backward
to near eye between the orbital bristles; face with similarly coloured streaks on
parafacial sutures, across lower margin, and on each side of central line of face,
the two latter connected above and below middle; antennae testaceous yellow;
aristae and palpi fuscous, the latter pale at tips. Thorax brownish testaceous,
slightly shining, with patches of dense whitish grey dusting on disc and pleura,
those on disc outlining two broad submedian vittae and a number of connected
markings laterad of them; scutellum dark brown, with a spot of whitish dust*in
front of base of each basal bristle. Abdomen shining testaceous, darker at bases
of the tergites. Legs testaceous, fore femora largely brown, mid and hind pairs
and all tibiae brown near bases and apices, third and fourth segments of fore
tarsus fuscous. Wings yellowish hyaline, fuscous from bases to apices of basal
cells and beyond humeral vein. Knobs of halteres fuscous.

Frons subquadrate, all bristles strong, anterior orbitals close to anterior

Margin and not farther from eyes than posterior pair; antennae normal; arista
5

416 NOTES ON AUSTRALIAN DIPTERA, Xiii,

pubescent; face flat. Thorax with 1+3 pairs of strong dorsocentrals, two intra-
dorsocentral series of short closely placed hairs, a pair of strong prescutellar
acrostichals, and two strong sternopleurals. Fore femur without a distinct comb;
all tibiae with preapical dorsal bristle, mid pair with one apical ventral bristle.
Inner cross-vein at about one-third from apex of discal cell; last section of fourth
vein three times as long as penultimate section.

Length, 4-4-5 mm.

Type and one paratype, Donnybrook, W.A., 29th Aug., 1926 (E. W. Ferguson).

Belongs to the same group as plumiseta and peterseni, but closer to suffusa,
the arista being pubescent, and the intradorsocentral hairs short and fine. It is
distinguished from sujffusa by the presence of fuscous markings on the wings
being confined to the bases, and by the thoracic markings.

SAPROMYZA OCELLARIS, n. sp. (Text-fig. 3.)

Male and female.—Head testaceous, frontal orbits, face, and cheeks, densely
white dusted, interfrontalia yellow, ocellar spot, a line along inner margin of each
orbital stripe, and a triangular mark between each antenna and eye, velvety black;
face with a pair of brown spots at middle; antennae, aristae, and palpi, black;
occiput with a dark mark behind upper angle of each eye, and a brown streak on
lower half. Thorax dull brownish, or testaceous yellow, densely grey dusted,
dorsum with four dark brown vittae, the submedian pair fused behind and carried
on to disc, but not to apex, of scutellum; pleura with the following dark brown
marks; a streak on upper margin of mesopleura, another on its lower margin,
a third one below sternopleural bristles, and two spots on centre of mesopleura.
Abdomen coloured as thorax, tergites with a dark central vitta, and dark spots at
bases of the larger bristles. Legs testaceous, femora sometimes brownish or
fuscous, fore legs from near bases of tibiae, apices of mid and hind tibiae and tarsi
black. Wings with many dark brown markings, many of them ocellate (Text-
fig. 3). Halteres yellow.

Frons a little longer than wide, all bristles long, orbits broad and distinct;
face in profile quite prominent above, with a transverse impression below; third
antennal segment about twice as long as wide; arista with rather dense hairs,
the longest less than half as long as width of third antennal segment. Thorax
with three pairs of dorsocentral bristles, the anterior pair close to suture, the
intradorsocentral hairs in four series, the inner two rather long,:the outer two
irregular, and the prescutellar acrostichal pair long; scutellum flat, rather long.
Fore femur with a very weak anteroventral comb; mid tibia with three apical
ventral bristles, one much weaker than the others.

Length, 3-4 mm.

Type, male, allotype, and one male and one female paratypes, Burnie, Tasmania,
24th and 27th Oct., 1922; paratypes, two, Adventure Bay, Tasmania, 27th Dec.,
1922, and ist Jan., 1923; one, Geeveston, Tasmania, 8th Dec., 1922; one, Hagle-
hawk Neck, Tasmania, 3rd Nov., 1922; one St. Patrick R., Tasmania, 30th Oct.,
1922; one, Adelaide, S.A., 20th Oct., 1921 (A. L. Tonnoir).

This species runs to section 20 in my published key to the species of this
genus, but is distinguished from any species in the subsequent part of the key,
and in fact from any other Australian species, by the wing markings.

SAPROMYZA AVICOLA, N. SD.
Male.—Belongs to the group in which regalis Malloch occurs, running to this
species in my published key. It differs from regalis in having the humeri normal,

BY J. R. MALLOCH. 417

not exceptionally prominent, with one long bristle and a few very weak hairs, and
the hairs on adjoining region sparse and fine close to humerus, becoming more
numerous and stouter beyond, but not so conspicuous as in regalis. The post-
humeral bristle is quite short. Inner cross-vein at a point about one-fourth from
apex of second costal division. Mid tibia with one apical ventral bristle.
Length, 6 mm.
Type, Eaglehawk Neck, Tasmania, 18th Nov., 1922 (A. L. Tonnoir).

SAPROMYZA TONNOIRI, Nl. SP.

Male.—Glossy fulvous yellow, entire frons shining; ocellar spot, two sub-
triangular spots near centre of face, and usually one near each vibrissal region,
fuscous; antennae and palpi yellow; arista fuscous. Abdomen glossy black on
dorsum except the basal tergite. Legs and halteres yellow. Wings hyaline.

Frons broader than long, slightly convex, all bristles present, anterior orbitals
much weaker than the posterior pair; face slightly convex; arista pubescent.
Thorax with the dorsocentrals arranged 1+3, and usually a similar number of
acrostichals, the latter rather irregular and not strong; sternopleurals 2; scutellum
rather short and convex. Fore femur without an anteroventral comb; all tibiae
with a preapical dorsal bristle, mid tibia with one apical ventral bristle. Inner
cross-vein much beyond level of apex of first vein, and about two-fifths from apex
of discal cell; ultimate section of fourth vein about twice as long as penultimate
section.

Length, 2:75-3:25 mm.

Type, Launceston, Tasmania, 9th Oct., 1922; single paratypes, Wilmot,
Tasmania, 8th Jan., 1923; Hobart, Tasmania, 4th Jan., 1923; and Zeehan, Tasmania,
7th Feb., 1923 (A. L. Tonnoir).

This species runs to section 5 in my published key to the species of this genus,
but is distinguished from any species between that section and 8, in which all
those with four pairs of dorsocentral bristles and unmarked wings are included,
by the marked face, fulvous yellow thorax, and pubescent arista.

SAPROMYZA STRAHANT, N. SD.

Male.—Fulvous yellow, thorax and abdomen glossy. Frons with the orbital
stripes a little more shining than remainder of frons, ocellar spot fuscous; face
shining, whitish dusted on sides; antennae and aristae black, third segment of
former a little reddish at base; palpi yellow, darkened at apices. Thorax with a
faint dark streak along upper margin of mesopleura. Abdominal tergites in type
specimen slightly dark at apices. Legs black, all coxae, mid and hind femora
except their apices, bases of mid tibiae, and basal segment of fore and hind tarsi,
yellow. Wings slightly and almost evenly smoky, darkest along costa. MHalteres
yellow.

Frons a little wider than long, orbits slightly differentiated; ocellar bristles
microscopic; anterior pair of orbitals about half as long as posterior pair, much
closer together than these, distance between them not much greater than distance
of either from eye; third antennal segment slightly tapered, about twice as long
as its basal width; arista microscopically pubescent; face almost flat, and a little
receding below. Thorax with three pairs of strong dorsocentrals, the anterior
pair close to suture, intradorsocentral hairs very short, in four series in front
only, and the inner postalar and anterior sternopleural bristles weak. Fore femur
with an anteroventral comb; ali tibiae with a preapical dorsal bristle; mid pair

418 NOTES ON AUSTRALIAN DIPTERA, Xiii, *

with one long apical ventral bristle. Inner cross-vein slightly beyond middle of .
discal cell and level of apex of first vein; ultimate section of fourth vein not over
1-5 as long as penultimate section. 5

Length, 4 mm.

Type, Strahan, Tasmania, 6th Feb., 1923 (A. L. Tonnoir).

This species runs to section 23 in my key but is distinguished from any in the
subsequent part of key by the fulvous yellow thorax, which is not dusted nor

vittate.

7

SAPROMYZA FLAVODORSALIS, D. Sp.

Male and female.—Head testaceous yellow; frons orange-yellow, shining only
on the orbital stripes, the latter, vertex, and ocellar region, greyish dusted; ocellar
spot and upper occiput fuscous, grey dusted; antennae and aristae black, basal
antennal segment rufous; palpi black. Thorax fuscous, dorsum, including the
humeral angles, fulvous testaceous, the whole greyish dusted, dorsum shining.
Abdomen fuscous, slightly shining, with dense even grey dust. Legs testaceous
yellow, blackened on all of fore femora and bases of mid and hind pairs, all except
bases of fore tibiae, and the apices of mid and hind pairs, all of fore tarsi and the
apices of mid and hind pairs. Wings yellowish hyaline. Halteres yellow.

Frons not longer than wide, with some microscopic black surface hairs, all
bristles distinct; anterior orbitals about one-third from anterior margin of frons
and farther from eyes than posterior pair; third antennal segment rather wide at
base, tapered to apex, and not twice as long as its basal width; arista practically
bare; face concave in profile, a little receding below; cheek not as high as width of
third antennal segment. Thorax with three pairs of postsutural dorsocentral
bristles, the anterior pair quite short and rather close to suture, the hairs anterior
to the bristles quite long and strong so that sometimes there may be four, or even
more, almost bristle-like; intradorsocentral hairs biseriate, two or three of the
pairs behind, and sometimes even up to or in front of suture, quite long, and
laterad of these two series there are usually some scattered short hairs so that in
some cases the hairs may appear quadriseriate; sternopleurals two. Fore femur
without an anteroventral comb; all tibiae with distinct preapical dorsal bristle,
mid pair. with one long and one very short apical ventral bristle. Wings narrow;
inner cross-vein very slightly before the middle of discal cell and distinctly before
apex of first vein; ultimate section of fourth vein not over 1:5 as long as
penultimate section.

Length, 4 mm.

Type, male, and allotype, Mt. Field, Tasmania, 21st Dec., 1922 (A. L. Tonnoir).

If this species is accepted as having the anterior pair of postsutural dorso-
centrals much weaker than the posterior two pairs, which is evidently the case,
it will run to section 11 in my key, but it differs from any species in the sections
between 11 and 19 inclusive in having the mesonotum fulvous testaceous and
the pleura and metanotum fuscous. If it is placed amongst the species with three
strong pairs of postsutural dorsocentrals it will run to section 23, but, like
strahani, it differs from any included in the subsequent sections in the colour
of the thorax.

15. Genus TRYPANEOIDES Tonnoir and Malloch.

This genus contains, so far as I know at present, species which are as a rule a
little smaller than the average of those in Sapromyza, some being less than 2 mm.

BY J. R. MALLOCH. 419

in length, and the wings of the known species are always conspicuously pictured.
The presence of a strong bristle on middle of the mesopleura, which is always
more or less downwardly directed, readily distinguishes the genus from any other
except one which occurs in Formosa. : The species very closely resemble small
Trypetidae. There are no species of the genus known to me from Australia, the
genotype was described from New Zealand, but there are about a dozen species
distributed throughout the Orient.

16. Genus HomonrurRA van der Wulp.

I have in the press a paper in which I erect some new subgenera of this genus
and two of the species now before me from Australia are used as the types of new
subgenera in this paper.

HoMONEURA (EUHOMONEURA) ORNATIPENNIS (de Meijere).

I have compared the type specimen of this species with that of atrogrisec
Malloch and can find no characters that would justify me in distinguishing them as
different species. Originally the species was described from the Nederland Indies
and its occurrence in Australia may be due to commerce, though there is nothing
yet known of its larval and pupal stages to indicate how it may be transported.

I erect for this species and three others which occur in the Orient the new
subgenus, Euhomoneura, with ornatipennis as type.

The distinguishing character is found in the dorsocentral bristles of thorax
which are arranged 1+2. The most closely related subgenus is Xenohomoneura
described below and the differentiating characters are noted thereunder.

Subgenus XENOHOMONEURA novum.

This subgenus is erected for a species which has the same arrangement of the
dorsocentral bristles of thorax as has Ewhomoneura, but the wings are hyaline,
not conspicuously pictured as in that subgenus, and the distance from lateral
edge of central portion of face to eye and the width of central portion and its lower
angle are about equal. In Euhomoneura the central portion of face is about three
times as wide at lower margin as the distance from its edge to eye. The type is
testaced, n. sp.

HoMONEURA (XENOHOMONEURA) TESTACEA, 1. SD.

Male and female.—Dull testaceous yellow; aristae and ocellar spot fuscous;
dise of mesonotum, and to a lesser extent that of scutellum, infuscated, and rather
copiously grey dusted; metanotum fuscous, and grey dusted; abdomen of female
with a dark dorsocentral vitta. Wings hyaline.

Frons about one-third of the head width and a little longer than wide, all the
bristles strong, the anterior orbitals close to anterior margin of frons; antennae
rather small, third segment about 1-5 as long as wide; arista minutely pubescent;
lower occiput slightly bulging; cheek higher than width of third antennal
segment; face almost flat. Thorax with all three pairs of dorsocentrals long and
strong, the prescutellar acrostichals short and fine, and the intradorsocentral hairs
in four irregular series; scutellum flattened above, not elongated; sternopleurals
both present. Fore femoral comb not conspicuous; all tibiae with preapical dorsal
bristle; mid pair with two apical ventral bristles. Inner cross-vein at about two-

420 NOTES ON AUSTRALIAN DIPTERA, Xiii,

fifths from apex of discal cell; ultimate section of fourth vein less than twice as
long as penultimate section.

Length, 3 mm.

Type, male, and allotype, Adventure Bay, Tasmania, 7th Dec., 1922 (A. L.
Tonnoir).

HoMONEURA (HOMONEURA) FERGUSONI, Nn. SD.

Male and female.—Testaceous yellow, slightly shining. Antennae and palpi
yellow; aristae fuscous. Thorax slightly dusted, usually with faint traces of two
brownish vittae just mesiad of the dorsocentrals. Abdomen of male usually with
a faint series of dark dorsocentral spots on apical half, that of female with the
spots quite distinct and in addition a larger one on each side of the fifth tergite.
Legs yellow. Wings hyaline, normally with seven dark spots, those over inner
cross-vein and at least the anterior extremity of outer cross-vein, just before apex
of second vein, and at' apex of third vein, most distinct, those at apex of fourth
vein, at base of antepenultimate section of same vein, and between apices of
auxiliary and first veins, least noticeable, sometimes very faint. Halteres yellow.

Frons subquadrate, orbits faintly differentiated, all the bristles long, surface
hairs microscopic and sparse; third antennal segment almost twice as long as
wide, slightly angulate on upper apical extremity; arista with very short
pubescence. Thorax with three pairs of postsutural dorsocentrals and six series
of intradorsocentral setulae. Fore femur with an anteroventral comb; all tibiae
with distinct preapical dorsal bristle;; mid tibia with two subequal moderately
long apical ventral bristles. Inner cross-vein about two-fifths from apex of discal
cell; ultimate section of fourth vein a little longer than penultimate section.

Length, 4-5 mm.

Type, male, allotype and four paratypes, Geraldton, W.A., 5th Sept., 1926
(EK. W. Ferguson).

_ Named in honour of the collector, to whom I am indebted for most of the
material received from this region.

In my recently published key (These ProcrEpiNnGs, 1927, p. 12) to the species
of this genus from Australia this species will run down to section 4, but is
distinguished from apicinebula Malloch, the only species in the first alternative,:
by the fact that the dark spot on second vein is not at extreme apex, and the base
of antepenultimate section of fourth vein is clouded; from the second alternative
it is distinguished by the pubescent arista.

HoMONEURA (HOMONEURA) FLAVOFEMORATA, TD. Sp.

Male and female.—Fulvous yellow, thorax and abdomen glossy, frons except
the orbits, sides of face, and cheeks, not shining. Ocellar region with a dark
mark on inner margin of each ocellus; antennae yellow, third segment and aristae,
and the apices of palpi, black. Legs yellow, blackened from near bases of tibiae
to apices of tarsi. Wings yellowish hyaline, both cross-veins slightly clouded.
Halteres yellow.

Frons subquadrate, all bristles long, anterior orbitals a little farther from
eye than posterior pair and about one-third from anterior margin of frons;
third antennal segment about 1:5 as long as wide; arista microscopically
pubescent; face slightly convex in profile; cheek fully as high as width of third
antennal segment. Thorax with three pairs of strong postsutural dorsocentrals,
the anterior pair close to suture, and eight or more irregular series of intra-

BY J. R. MALLOCH. 421

dorsocentral hairs; scutellum slightly flattened. Female with about eight short
bristles on genital segment. Fore femur with a slight anteroventral comb; all
tibiae with a distinct preapical dorsal bristle, mid pair with two apical ventral
bristles. Inner cross-vein a little beyond apex of first vein and middle of discal cell;
ultimate section of fourth vein not 1:5 as long as penultimate section.

Length, 4-4-5 mm.

Type, male, and allotype, Haglehawk Neck, Tasmania, 15th Nov., 1922
(A. L. Tonnoir).

This species runs down to section 12 in my key to the species of this genus,
but is distinguished from all the species included thereinafter by the colour of
the legs, all the others having the tibiae, and usually also the tarsi, yellow.

17. Genus AusTRALINA, Malloch.

I have seen only the two original specimens of the genotype of: this genus,
geniseta Malloch, described from Darwin, N.T.

There are some species of this family still in my hands which I do not care
to describe from single specimens in rather teneral condition.

Family Agromyzidae.
Genus CRYPTOCHAETUM Rondani.

This genus is of considerable economic importance in Australia and the
south-western United States, feeding upon certain scale insects, in which the
larvae are internal parasites. Two species were brought from Australia to
California about 35 years ago, and are now established there as parasites of a
scale insect infesting citrus trees. The genus occurs also in Hurope and Asia.

Two subgenera have been recognized by Bezzi, based upon the structure of
the frons, presence or absence of hairs on the eyes to some extent, and some
trivial wing characters. The known Australian species fall into the subgenus
Lestophonus, all having the eyes hairy, and the frontal triangle wide at anterior
margin. Below I give a synopsis of the characters by means of which the
Australian species Known to me may be recognized. I am accepting the specific
names applied to the two old species, but whether these are correct or not I cannot
be absolutely certain.

The genus is the only one in the Acalyptratae in which the antennal arista
is lacking (Text-fig. 4). The species have a habitus very similar to those of
Leucopis Meigen, which are so far as known predacious enemies of aphides and
scale-insects, and with which I linked them in a paper on Agromyzidae published
a number of years ago. The most recent work on the families of Acalyptratae
by Hendel has, however, placed a great deal of emphasis upon the structure of
the basal half of the costal vein of the wing and the degree of development and
individuality of the auxiliary vein, and if we accept these characters as criteria
for the segregation of families then Cryptochaetum can hardly be considered as
Ochthiphiline. All the members of this latter group have the auxiliary vein
complete, though sometimes closely approaching first vein near its apex, and the
costal vein is not broken at the point where the auxiliary vein connects with
it. In Cryptochaetum the auxiliary vein is almost obsolete, being visible only
under a very high power lens, and the costa is slightly interrupted where the
rudimentary auxiliary vein connects with it. There is no cross-vein between the
discal cell and the anterior basal one in Cryptochaetum, while there is always
such a cross-vein in the Ochthiphilinae.

422 NOTES ON AUSTRALIAN DIPTERA, Xiii,

‘All the species of the genus are bluish black in colour, and the- frontal
triangle, which is the same colour, is very large, occupying most of the frons,
the latter lacks distinct lateral bristles and, like the dorsum of thorax, scutellum,
mesopleura, and abdomen, is furnished with short hairs. Differences in venation,
and the structure of the fore tarsi in the males, are the principal Wee
characters.

Key to the Species.

1. Inner cross-vein of wing nearly one-third from apex of discal cell; ultimate section of
fifth vein not over two-thirds as long as penultimate section; third antennal
segment with a*sharp apical upper angle; fore tarsus in male without exceptional
Structures on shairs-mcalyptrae fuSCOuUS| aha e orien one monophlebi Skuse.

Inner cross-vein of wing close to, or even proximad of middle of discal cell;
ultimate section of fifth vein at least three-fourths as long as penultimate
section; third antennal segment rounded at apex, or obtuse. .............. 2

Fore tarsus of male dilated (Text-fig. 5); calyptrae white; inner cross-vein a little
beyond middle of discal cell; ultimate section of fifth vein about three-fourths
as plongyvas= penultimate! Section once serene eee eee latimana., Nn. sp.

Fore tarsus of male not dilated, with some short fine erect hairs along the anterior
side (Text-fig. 6); inner cross-vein sometimes a little proximad of middle of
discal cell; ultimate section of sea vein about as long as penultimate section
Feu EEE. SRCRCLORE EER CRORES He SET IIS TH eR ETE AL CRD aheome Om Le Ae ae ure tea iceryae Williston.

to

Text-fig. 4. Cryptochaetum latimana, head from side.
Text-fig. 5. Cryptochaetum latimana, fore tarsus of male.
Text-fig. 6.° Cryptochaetum iceryae, fore tarsus of male.

CRYPTOCHAETUM LATIMANA, Nn. Sp. (Text-figs. 4 and 5.)

Male.—Glossy blue-black, frons on sides of the triangle, and the antennae, deep
opaque black. Legs shining black, tarsi testaceous. Wings hyaline. Calyptrae
white. Halteres black.

Frontal triangle gradually narrowed from vertex to anterior margin, at the
latter point narrower than distance of its side from eye; third antennal segment
not angulate but broadly rounded at apex. Scutellum margined as in other
species. Abdomen rather wide, third and fourth visible povetLes subequal in length.
Fore tarsus as in Text-fig. 5.

Inner cross-vein almost exactly at middle of discal cell; penultimate section
of fourth vein a little over one-fourth as long as ultimate, the latter diverging from
third, and at its apex a little farther from third, measured on costa, than third is
from second; ultimate section of fifth vein about three-fourths as long as
penultimate.

Length, 1-5 mm.

Type, Wahroonga, Sydney, N.S.W., 24th Oct., 1926.

CRYPTOCHAETUM MONOPHLEBI Skuse.
I have before me Australian specimens of this species, or at least specimens
so labelled, and which agree with the characters cited for its distinction. Some

BY J. R. MALLOCH. 425

of the specimens were reared from’ Monophlebus. These specimens all have the
third antennal segment with a very pronounced angular production on its upper
apical angle which character appears to ally them very closely with curtipenne
Knab, the type specimen of which I have compared with the specimens referred
to. The type of Knab’s species is in very poor condition and it is impossible to
say whether or not it is the same species, though one of the Australian specimens
of monophlebi bears this name lahel. The male has the fore tarsus normal

CRYPTOCHAETUM ICERYAE Williston. (Text-fig. 6.)

Readily distinguished from the other two species by the characters cited in
the foregoing key. Fore tarsus of male as in Text-fig. 6. I have before me
. Australian and American specimens. .
There are probably Australian species of this genus yet undiscovered.

Genus CERopoNTA Rondani.

I have previously described two species of this genus from Australia and now
present a third one. To facilitate the identification of these three species I am
giving below a synopsis of their distinguishing characters.

Key to the Species:

1. Scutellum lemon yellow, with a deep black spot on each basal angle; thorax shining
yellow, mesonotum with the following glossy black marks; a broad complete
central vitta, a slightly narrower vitta each side of it, which does not extend
to anterior margin, and a narrower one behind suture laterad of these; humeri
with a black spot; intradorsocentral hairs in about six irregular series; third
antennal segment as in robusta, shorter than usual and acute at apex ........
fic ce to ph od 6-0 OD Op OR GOD Oe Ul cieaRohaia DiGsnLuN GG UNOS hens oucye cAsieneuchss Suen LULELLG ET, ons w SD:

Scutellum black on entire disc; thorax entirely or almost entirely black above .. 2

Stout species; pleura largely lemon yellow; disc of mesonotum black, with slight
grey dusting, the margins lemon yellow, the hind margin narrowly so; intra-
dorsocentral hairs in at least four irregular series on almost the entire iength
of thorax; apices of abdominal tergites lemon yellow; third antennal segment
shorter than usual and with a sharp apex; metanotum small .. robusta Malloch.

Thorax black, with dense grey dust, and nowhere distinctly shining, mesonotum with
two faint brownish vittae posteriorly on the lines of dorsocentrals; intradcorso-
central hairs biseriate, sparse, and present only anteriorly; abdominal tergites
black; third antennal segment normal, more elongate than in robusta, and with
a sharp point at upper apical angle; metanotum prominent, extending to apex
OLE SCUCES LL Ure pave vere akey eeoc oie <hraioa ablative soieviodlc ua ey once oo artch ows tater hee palione move tenes australis Malloch.

to

CERODONTA VITTIGERA, Nl. SDP.

Male.—Head orange-yellow, frontal orbits paler, ocellar region with a large
black mark; aristae black; third antennal segment with a very small part of
apex black; inner mouth margin and occiput black. Thorax lemon yellow, dorsum
with five glossy black vittae as described in the key; pleura largely black, the
sclerites yellow on margins. Abdomen black, apices of tergites yellow. Wings
hyaline. Halteres yellow.

Head as in robusta. Thorax with 1 + 3 pairs of dorsocentrals, about six
series of long intradorsocentral hairs which are continued almost to scutellum,
the intra-alar bristle hardly distinguishable from the hairs, and no differentiated
prescutellar acrostichals; metanotum not well developed. Legs stout, the femora
quite pronouncedly so. Wings normal, inner cross-vein at about one-third from
apex of discal cell, outer cross-vein vestigial in both wings, ultimate section of

424 NOTES ON AUSTRALIAN DIPTERA, Xiii,

fourth vein about nine times as long as penultimate; ultimate section of fifth
vein longer than penultimate. ;

Length, 2:75 mm.

Type, Koolanooka, W.A., 9th Sept., 1926 (H. W. Ferguson).

Genus AGroMyza Fallen.

I have been accumulating material for a review of the Australian species of
this genus for some time, but have not yet obtained sufficient to warrant my
publishing on the entire genus. Below I deal with some of the available species
which belong to the group in which the halteres dre entirely black or fuscous,
and the entire insect is black, usually with some part of thorax or abdomen, or
both, with a distinct blue or green tinge, sometimes quite noticeably metallic.
There are at least two segregates of this group, those before me from Australia
being referable to Melanagromyza Hendel, and Ophiomyia Braschnikow. These
have been accepted by Hendel as genera, but I think subgeneric rank is all they
are entitled to, and I so treat them herein. Below I present a synopsis of the
characters that are of value in distinguishing them. Hendel has cited the
difference in the course of the auxiliary vein in the two major groups in his recent
paper on the European species of Agromyza as a character for their separation,
but I can not see any fundamental distinction here. In neither group is this vein
absolutely fused with the first vein according to my opinion, though in certain
species it appears to be so owing to the bending up of the membrane between the
first vein and the vestigial auxiliary vein. If the wing is carefully flattened out,
or turned round, in examining it it will be seen that there is practically no
distinction between the two groups that are so distinguished in his key.

Synopsis of Subgenera.

A. Face without a broad central vertical carina, sometimes with a linear one which is
not much elevated and does not appreciably separate the antennae at bases,
the antennae projecting parallel, or almost so; vibrissal angle not prominently
projecting and never with a fasciculus of bristles in male ................
BE SiC OAC CaoNaG chohce ciececiC iotiomacokcrGar coun aa eeaes Ghitict or pei CEG Dn thorck cvovarciea erosonG Melanagromyza Hendel.

AA. Face with a quite prominent central vertical carina, more or less widened above
middle, which very distinctly separates bases of antennae, the antennae almost
invariably distinctly diverging to sides; vibrissal angle projecting, and with
two or more long bristles forming a curved fasciculus in male ..............
ite ett tte ape taae Melis, oh ainda fondee Wi ena aa le ahah er leb antes tae kd SRI Ophiomyia Braschnikow.

Subgenus MELANAGROMYZA Hendel.

The larvae of the great majority of the species in this subgenus mine in the
roots and stems of low plants and some are quite destructive. One of the worst
species is M. phaseoli Coquillett, which affects beans in Australia.

In the key given below I include three named and two unnamed species.

Key to the Species.

1. Thorax with three pairs of well developed dorsocentral bristles .... trispina, n. sp.
Thorax with but two pairs of distinct dorsocentral bristles .................-.+.+- 2
2. Calyptrae white, their margins yellowish, or whitish, and fringes white ........ 3
Calyptrae greyish or fuscous, their margins and fringes fuscous, or dark brown .. 4

Inner cross-vein of wing at middle of discal cell; ultimate section of fourth vein
about three times as long as penultimate; ultimate section of fifth vein hardly
over half as long as penultimate ‘section 2 icici c cies. os albisquama, Nn. sp.

Inner cross-vein of wing about one-third from apex of discal cell; ultimate section
of fourth vein not less than five times as long as penultimate; ultimate section
of fifth vein about two-thirds as long as penultimate section ............ sp. 1

i)

BY J. R. MALLOCH. 425

4. Frontal triangle polished greenish black, extending narrowly to, or almost to, the

ENUekoye ions Ge IRIN ohovocaoocdondnoDeooU On HDOdNOGDOT phaseoli Coquillett.
Frontal triangle slightly shining, black, wider than in phaseoli, and extending to
NO UKE TMCS Oi OU 5 Gowecoes Joon ocbaddooodoaccdodoUoaoMddoUccaob obo DOM sp. 2

AGROMYZA (MELANAGROMYZA) TRISPINA, TD. SD.

Male.—Shining black. Frons shining on orbits and posterior part of triangle.
Seutellum slightly dusted and not so shiny as mesonotum. Legs black. Wings
hyaline. Calyptrae and their fringes white. Knobs of halteres black.

Eyes bare; frons over one-third of the head width; orbits narrow, each with
about six bristles, and many short hairs; triangle distinct only on posterior half,
obsolete anteriorly, its apex at about one-third from anterior margin of frons;
ocellar bristles erect and divergent, as long as postvertical pair; face slightly
carinate; vibrissal angle not produced, with a fine bristle and some long hairs;
antennae small, third segment rounded; arista thickened basally, subnude; cheek
barely as high as width of third antennal segment. Thorax with three pairs of
dorsocentral bristles, the anterior pair weakest and distinctly behind the suture,
the intradorsocentral hairs in about ten series, prescutellar acrostichal bristles not
developed, postalar bristle next to scutellum minute; intra-alar bristle distinct.
Legs stout; mid tibia with one or two posterior setulae. Inner cross-vein at less
than one-third from apex of discal cell; outer cross-vein at a little less than its
own length from inner and over 1:5 its own length from apex of fifth vein;
ultimate section of latter over three-fourths as long as penultimate; ultimate
section of fourth vein deflected apically, and about five times as long as penultimate
section.

Length, 3 mm.

Type, Merredin, W.A., 31st Aug., 1926 (E. W. Ferguson).

It is exceptional for a species in this subgenus to have three pairs of dorso-
central bristles, nearly all having but two pairs.

AGROMYZA (MELANAGROMYZA) PHASEOLI Coquillett.

This species differs from the preceding one in having the dorsocentrals
consisting of two pairs, the fronto-orbital bristles usually five in number, with
the anterior two pairs incurved, the frontal triangle polished and extending
narrowly to anterior margin of frons, the face with a low sharp central vertical
carina, the vibrissal angle slightly produced and with one bristle, the inner cross-
vein at about two-fifths from apex of discal cell, the ultimate section of fourth
vein four or five times as long as penultimate, and the length of specimens about
1-5 mm.

A common enemy of beans.

AGROMYZA (MELANAGROMYZA) ALBISQUAMA, 0. SD.

Male.—More distinctly metallic green-black than phaseoli, and with the
calyptrae white, their margins yellowish and fringes white.

Structurally quite similar to phaseoli, but the frontal triangle is broader at
posterior margin and is more evenly narrowed from there to anterior extremity
which latter almost attains the anterior margin of frons. Thorax as in phaseoli.
Inner cross-vein at middle of discal cell; ultimate section of fourth vein three times
as long as penultimate; ultimate section of fifth vein about five-eighths as long as
penultimate section.

Length, 2 mm.

426 NOTES ON AUSTRALIAN DIPTERA, Xiii,

Type, Eccleston, Allyn R., N.S.W., 28th Feb., 1921.
In addition to the above I have two other species on had but it is not
advisable to deal with these on the basis of single specimens.

Subgenus OpHiomy14a Braschnikow, 1897.
There is but one Species of this subgenus before me at this time, unless a
single specimen mentioned hereinafter belongs to a distinct species.

AGROMYZA (OPHIOMYIA) LANTANAE Froggatt.

This species is the same length as phaseoli, but belongs to a different subgenus,
the face being more conspicuously carinate, with the carina broadened above
middle and distinctly separating the bases of antennae, and a more prominent
vibrissal angle on which there is a fasciculus of two or three long bristles in the
male, and at least one prominent bristle in female. In other respects the species
closely resemble each other, but the frontal triangle in 7Jantanae is less shiny,
and shorter and broader, extending only to about middle of frons. The inner
cross-vein is at, or close to, one-third from apex of discal cell, the ultimate section
of fourth vein is about seven times as long as penultimate section, and the
posterior bristles on mid tibiae are extremely short.

Feeds on Lantana. Narrandera, North Harbour, Mosman, and Sydney, N.S.W.

I have before me a female specimen in poor condition which belongs to this
subgenus, and may be a distinct species. It is a little larger than lantanae and
has the inner cross-vein very slightly beyond middle of the discal cell. Sydney,
N.S.W.

In connection with members of this genus, and especially those of the two
subgenera above dealt with, it is highly desirable that rearing records be obtained
wherever possible and that specimens of the mines, and larvae and puparia, be
kept when specimens are reared.

Subgenus Lraiomyza Mik.

This subgenus contains ‘species that are more slenderly built than in the two
preceding subgenera, which have the halteres always yellow, and the auxiliary
vein of the wing more evidently separated from the first at apex than appears to
be the case in these and in the subgenus Agromyza Fallen.

I note also that there is a more constant lack of an intra-alar bristle here
than in the other subgenera mentioned, but until I attempt a review of all the
subgenera occurring in Australia I do not care to indicate the importance of this
character. It appears to me at this time that the character used by Hendel for
the separation of Liriomyza from Dizgonomyza is too trivial for such a purpose
as it consists solely of a difference in the colour of the scutellum, yellow or partly
yellow in Liriomyza, entirely black, or at least without yellow, in Dizgonomyza.
The genotypes are not available to me so that it is impossible to determine if there
are other characters that may be used for their separation. I do not use
Dizgonomyza in my treatment of the Australian species at present.

I have several species of Liriomyza, as accepted by me, in my hands at this
time, one of which, pusilla Meigen, I have already recorded from Australia. This
species has a very wide range and feeds upon a great diversity of food-plants,
mining in their leaves. It has also a large number of names applied to it, some
of them at least being entitled to retention for subspecies and varieties. It may
be that with a fuller knowledge of the immature stages some of these subspecies

BY J. R. MALLOCH. 427

or varieties may be established as good species, but careful work will have to
be done to determine this point. I describe only the species which appear to me
to be perfectly distinct from any known to me from any part of the world.
Possibly some of the species recently described by Mr. M. Watt from New Zealand
may be found to occur in Australia, but it is imperative that very careful
examinations be made of any such species as appear to be identical with his as
there appears to be a group of similar appearing species in this part of the world
that only a specialist can definitely distinguish from each other.

AGRoMYZA (LIRIOMYZA) PALLIDICENTRALIS, TD. SD.

Female.—Head dull yellowish testaceous, upper occiput, ocellar spot, upper
half of third antennal segment, aristae and clypeus, blackened, face usually
with a dark mark in each fovea; frontal orbits and vertex each side of the ocellar
spot paler than the interfrontalia; palpi yellow. Thorax fuscous, humeri and a
stripe along centre of scutellum testaceous yellow, mesonotum brown dusted and
entirely dull, with slight traces of darker vittae along the dorsocentral ‘lines.
Abdomen black, almost entirely dull, slightly grey dusted, and without conspicuous
yellow apices to the tergites. Legs dusky testaceous, irregularly suffused with
fuscous. Wings greyish hyaline. Calyptrae whitish, fringes pale brown. Halteres
white.

Frons fully one-half of the head width, orbits narrow, well defined, each with
three bristles, two outwardly and backwardly directed on upper half and one
inwardly directed on anterior half, besides a short hair which is sometimes present
anterior to the latter; ocellars longer than the postverticals; frons slightly’
projecting in profile; face with a slight central vertical carina; antennae of
average size, third segment rounded at apex; arista bare; cheek at anterior
margin about as high as width of third antennal segment, much higher at
posterior margin, the marginal hairs very fine and sparse, vibrissa short and fine.
Thorax with four pairs of dorsocentral bristles (1 + 3), no prescutellar acrostichals
nor intra-alar, and almost devoid of surface hairs, at most with a few between
anterior dorsocentrals; scutellum with four equal bristles. Legs slender, fore
femora without posteroventral bristles; mid tibia without any posterior median
setulae. Inner cross-vein slightly before apex of first vein and at middle of discal
cell; ultimate section of fourth vein about five times as long as penultimate;
ultimate section of fifth vein about 1-25 as long as penultimate.

Length, 2 mm.

Type and two paratypes, Sydney, N.S.W., 10th Oct., 1923, 26th Sept., 1923, and
24th Sept., 1924.

This species has much in common with flavocentralis Watt, but the wing vena-
tion is quite different, flavocentralis having the ultimate section of fifth vein over
twice as long as penultimate, and there are other distinctions.

AGROMYZA (LIRIOMYZA) TRICOLOR, 0. SD.

Male and female.—Head clay-yellow, interfrontalia more orange-yellow; ocellar
spot, and upper occiput except on margins, and centre above, black; third antennal
segment brownish above; aristae fuscous; palpi yellow; clypeus narrowly black.
Thorax clay-yellow, mesonotum fuscous except between the lateral Margins, the
dark part densely grey dusted except between the dorsocentrals from near anterior
margin, the central part brown dusted; scutellum fuscous, yellow at apex, densely
grey dusted on sides, brown dusted in centre; metanotum broadly shining black

428 NOTES ON AUSTRALIAN DIPTERA, Xiii,

centrally; a black mark on each humerus, one behind it on notopleural suture,
one on lower margin of mesopleura, another on lower part of pteropleura, and a
large black triangle on ‘the sternopleura. Abdomen clay-yellow, more or less
brownish on centre of dorsum, the genitalia in both sexes black. Legs clay-
yellow. Wings hyaline. Halteres clay-yellow, with a large brownish spot on each
knob.

Frons fully one-third of the head width; orbits distinct, each with three
bristles; ocellar and postvertical bristles subequal; third antennal segment rather
large, rounded at apex; arista with short pubescence; face slightly carinate in
centre; cheek at anterior extremity about half as high as width of third antennal
segment, much higher posteriorly; vibrissae short. Thorax with four pairs of
strong dorsocentral bristles (1 + 3), four series of intradorsocentral hairs anteriorly
only, and no prescutellar acrostichal nor intra-alar bristles; scutellum slightly
elongated, and narrowed at apex, slightly free from postscutellum at apex. Legs
as in preceding species. Wing almost exactly as in Watt’s figure of that of
citreifemorata Watt, inner cross-vein slightly before middle of discal cell; ultimate
section of fifth vein about twice as long as penultimate; ultimate section of fourth
vein fully four times as long as penultimate.

Length, 1-5 mm.

Type male and allotype, Como, N.S.W., December, 1923, swept from flowers
(H. Petersen).

Distinguished from any species in Australasia by the tricoloured thoracic
dorsum.

Family Chloropidae.
Subfamily CHLOROPINAE.
Genus PAcCHYLOPHUS Loew.

I have already described one species of this genus from Australia, and now
introduce other two, one of them so closely allied to lutea Malloch that it is not
necessary to give a detailed description of it. The other species is rather divergent
from type and I have included the essential characters for its separation from the
other two in the following synoptic key. The genus has much the appearance
of Meromyza Meigen, which genus contains species which mine in the stems
of wheat and other cereal plants, causing what are known as ‘“dead-heads”, but
nothing is at present known of the larval habits of the species of Pachylophus.

Key to the Species.

1. Hind femora with weak pale spinules in two faint series on a portion of apical half
of ventral surface, which are evident only under a very high power lens; face
without a dark mark in each antennal fovea, but with a complete blackish line
along each parafacial suture; arista gradually tapered from near base of third
segment to its apex; notopleural bristles 1 +1 .................. alienus, n. sp.

Hind femora with quite evident minute black spines in two series on at least the
apical half of ventral surface; face with a dark mark in each antennal fovea
on upper portion, the parafacial suture not blackened; arista lanceolate, about
equally wide almost to apex; notopleural bristles 1 + 2 ...............0...0% 2

2. Upper inner mouth margih with a central dark mark; pleura with a black streak
along lower margin of mesopleura and a black spot in centre of hypopleura,
sternopleura with most of its surface red, the upper posterior angle of the red
mark faintly blackish, the dark mark below prothoracic spiracle very faint,
eater] 00 A oe A UNE On SU coer eos Meade Ae A ee A NS oa as yl ok din Reet a ne RD A lutea. Malloch,

Upper inner mouth margin entirely black; pleura with a black streak along lower
margin of mesopleura and another along upper margin of the red sternopleural
area, a black mark in centre of hypopleura, one below prothoracic spiracle, and
a poorly denned Mark on preropleuiraii. a6 + sikelele ciaiekel Seo eres secundus, n. Sp.

BY J. R. MALLOCH. . 429

PACHYLOPHUS SECUNDUS, 0. SD.

In addition to the characters cited in the above key this species is more
slender and a little smaller than lutea, and though both have the characteristic
black streaks on upper part of face and the black spot on each fore coxa at base,
these are more conspicuous here than in lutea. The three dorsal vittae on the
thorax are quite variable in colour, being sometimes nearly all rufous, and varying
to nearly all black.

Length, 2:5-3 mm.

Type and four paratypes, Coolgardie, W.A., 12th Sept., 1926.

PACHYLOPHUS ALIENUS, 0. Sp.

Female.—Shining testaceous yellow like the other two species, but the frontal
triangle has a rather broad black streak along each side, the parafacial suture has
a complete blackish line on it, there are two fuscous spots on clypeus, the central
mésonotal vitta is reddish, and the posterior two-thirds of the submedian, and all
of the sublateral vittae, are black. There are five blackish spots on pleura, the
one on sternopleura being confined to the upper margin of the red mark, and
there is a black spot on each humerus.

Structurally similar to lutea except as stated in the key, and in having the
hind femora less thickened.

Length, 3-75 mm.

Type, Woy Woy, N.S.W., November, 1921 (A. L. Tonnoir).

It is possible that the presence of but one bristle on hind part of the noto-
pleura is abnormal, if not it is an interesting digression from type.

Genus CHLoRoPIScCA Loew.

This genus is the only one in this subfamily known to me which has a distinct
oval sensory area on dorsal surface of the hind tibia. This sensory area consists
of an elongate flattened section covering a variable extent of the posterodorsal
surface of the tibia, generally slightly darker in colour than the remainder of the
surface, and is furnished with microscopic erect pile. When I first mentioned
the existence of this area in the literature of this family I considered it possible
that it was of a sensory nature, and suggested that some morphologist examine
it to discover its function if any. So far as I know this has not yet been done.
The character is present in nearly all genera in Botanobiinae, but I have not
noticed it in related families, though there exists on the basal segment of the fore
tarsus of males of the South American Sapromyzid genus Griphoneura a small
flattened area that suggests a similarity to the one under consideration, though
here there are no erect hairs present.

I present below the description of a species from Australia which appears to
be undescribed, though it closely resembles notata Meigen.

CHLOROPISCA SUBNOTATA, n. Sp. (Text-fig. 7.)

Female.—Head yellow, ocellar spot, apical margin of third antennal segment
and the entire arista, black; frontal triangle more or less browned, but nowhere
black; inner mouth margin sometimes brownish. Thorax with the usual five
glossy black vittae, submedian pair not extending to anterior margin, laterals not
extending in front of suture; pleura with a black spot on mesopleura and another
on hypopleura, sternopleura red below, sometimes with a black spot above;
metanotum black in centre; scutellum yellow. Abdomen with most of the tergites

430 NOTES ON AUSTRALIAN DIPTERA, Xiii,

blackened, fourth with two. black spots. . Legs yellow, fore tarsi slightly dark at
apices. Wings hyaline. Halteres yellow. . :
Frontal triangle almost as wide as vertex, impunctate, with numerous fine
lateral hairs, sides almost straight, apex almost at anterior margin of frons; arista
subnude; cheek about as high as width of third antennal segment. Dorsum of

Text-fig. 7. Chloropisca subnotata, wing.
Text-fig. 8. Chloropisca monticola, wing.
Text-fig. 9. Diplotoxa tasmaniensis, wing.

thorax with numerous fine short hairs; scutellum flat, elongate, with two long and
two very short hairs. Fore tarsus slightly dilated apically. Wings as in Text-fig. 7,
subcostal cell very narrow.

Length, 2-5 mm.

Type and paratype, Mosman, N.S.W., 2nd Apr., 1923.

Resembles notata Meigen, but it has the frontal triangle quite coarsely punctate
on sides.

CHLOROPISCA MONTICOLA, n. Sp. (Text-fig. 8.)

Female.—Head testaceous, upper occiput and entire frontal triangle glossy
black; third antennal segment browned above; aristae fuscous; inner mouth margin
hardly darkened; palpi yellow. Mesonotum glossy black, more brownish anteriorly
on sides, yellow on lateral margins; scutellum yellow; metanotum black; pleura
with four large black marks, situated on mesopleura, sternopleura, pteropleura,
and hypopleura. Abdomen largely blackish brown above. Legs yellow, fore tarsi
hardly darkened apically. Wings hyaline. MHalteres yellow.

Frons a little over one-third of the head width, triangle not filling entire
vertex, its sides slightly curved, apex extending almost to anterior margin of frons,
surface smooth and polished; antennae normal; cheek about half as high as width
of third antennal segment. Thoracic dorsum smooth, the hairs rather numerous
and long; scutellum flat above, without a pronounced marginal rim, not elongate,
the disc with a few black hairs, and apex bristled as in preceding species. Fore
tarsus not widened, Wing as in Text-fig. 8.

Length, 1-75 mm.

Type, Blue Mts., N.S.W., 26th Mar., 1923.

This species is readily distinguished from suwbnotata by the entirely black
frontal triangle, black disc of mesonotum, and different venation of the wings. |

BY J..R. MALLOCH. 431

Genus CHLOROMERUS Becker.

This genus is readily distinguished from its allies by the thickened hind femora
with the short dense spines on the apical halves of their ventral surfaces, and
the normal venation, and shape of third antennal segment.

There are two described species, both from Australia, and to these I now add
four new species. It is particularly difficult for the taxonomist to determine from
a few dry specimens of certain genera in this family what are the specific limits.
Some species vary very much in colour and markings and without large series it is
often impossible to make certain of the specific characters. At times the colours
and markings are very stable, but in other cases so variable that only careful field
observations and critical examinations of series will suffice to identify species.

This condition of apparent variability obtains in this genus and it is impossible
for me to be certain of whether the forms of purus described herein are really
forms or valid species on the basis of the material on hand. It is quite possible
that they will turn out to be distinct species, but a larger series of each is a
desideratum to aid in determining this point.

Key to the Species.

1. All femora quite conspicuously marked with black ............... 0.00. c cece eee eee 2
Femora entirely yellow, or only the fore and mid pairs with a faint blackish ventral
TIMP TNE LS mates a scuch una uals ce Lauete Lancet sac cheed Aas 2) MRP en A sce a ARAB a cadens) pane te ee 3

2. Femora almost entirely glossy black; thoracic vittae and pleural spots entirely black;
clypeus (inner mouth margin) fuscous ..............00-2000- nigrifemur, nN. sp.
Femora yellow, with a blackish mark on upper surface at middle of each; thoracic
vittae largely red, central one with a pair of small black spots on hind extremity,
submedian pair each with a mark on outer side in front of suture, and the apical

half, black, sublateral pair entirely black and present behind suture only;
sternopleural mark red below, black on upper margin; clypeus yellow

aL cpr ON hi CG ha Pate tchles Pea cept ee Ra re Pla eel ely eT RY maculifemur, n. sp.

38. Slender species, with the hind femora not much thickened, not over twice as thick
as mid pair; frontal triangle entirely yellow, and without distinct lateral impressed

lines; third antennal segment almost wholly black ............ gracilis, n. sp.
Robust species, with hind femora much thickened, at least three times as thick as mid
pair; frontal triangle either with an impressed line along each lateral margin

which is usually quite conspicuously blackened, or the third antennal segment is

entirely yellow or slightly blackened on its upper margin .................. 4

4. Frontal triangle yellow, only the ocellar spot black; mesopleura, sternopleura, and
hypopleunassecachwwith: anblack mark wesimemeicrscieletondcncrckeiioie eee oeicleie oe rai 5
Frontal triangle yellow, with the lateral margins and ocellar spot black; pleura
usuallymwithyfives black: Marks) Vyas tects ro eleleysiel syeie elelielclsle) eel leu: (purus Becker.) 6

5. Clypeus yellow; thoracic dorsum with three glossy black vittae, due to a fusion of the
sublateral and submedian pairs behind suture ................ pallidior, Becker.

Clypeus partly black; thoracic dorsum with five glossy black vittae

5 6:0 0.010.) 0:0, 0'6'b,010:0 16 .Bis WBS ciel Old Bia/oto lors Ercole GiOlGuULD oir o Brawl lata tO O.oIORO.G trimaculata, n. sp.

6. Thoracic vittae largely red, only the posterior extremities, and sometimes the
sublateral pair on outer edges anterior to suture, black

NRTA Coe RAs sivsh si ausierasan vou ay lehnyanchrath miisianlar al etehaiis feuay silence purus, form varians, novum.
Thoracic vittae entirely or almost entirely glossy black ...................... 7
Thoracic vittae totally glossy black; fore and mid femora each with a faint, but
evident, blackish spot on ventral surface at middle ........................---
BS oIBIS o-d Ato laoid Gaol at EI OIG RIG IOIOI OS DIG CLO CG Ooi CMOS Ono purus, form maculifera, novum.
Median thoracic vitta usually reddish near anterior extremity; fore and mid femora

ENbirelivseviellOwy Veve cate! Sis ao sisters Sretaciele ta slit etait tile, euelene deus purus, form purus, Becker.

SH

CHLOROMERUS NIGRIFEMUR, 0. SP.
Male and female.—Head yellow, face more whitish, upper central part of
occiput and especially a line on each side at angles of frontal triangle darkened;

ocellar spot and a line along each side of triangle black; aristae fuscous; antennae
G

432 NOTES ON AUSTRALIAN DIPTERA, Xiii,

yellow, third segment darkened at insertion of arista; palpi yellow; clypeus
fuscous. Thorax with three broad black dorsal vittae, and an elongate black mark
on each side of these behind suture; humeri with a black spot; pleura with the
usual five black marks, the one on sternopleura covering all except its upper
margin. Abdomen brownish yellow, the apices of tergites paler, sometimes with
two dark spots on each side of first tergite. Legs yellow, femora almost entirely
glossy black. Wings hyaline. Halteres pale yellow.

Frontal triangle extending to anterior margin of frons, with an impressed line
along each side; ocellar bristles divergent, forwardly directed; antennae short,
distinctly separated at bases, third segment orbicular; arista microscopically
pubescent, thickened basally, second segment about three times as long as thick;
face not carinate; proboscis short; palpi normal; cheek about half as high as eye.
Dorsum of thorax slightly punctured at bases of hairs. Not much difference
between the three principal sections of costa.

Length, 3-3-25 mm.

Type, female allotype, and two female paratypes, Cradle Valley, Tasmania,
16th December, 1923 (A. L. Tonnoir).

CHLOROMERUS MACULIFEMUR, N. SDP.

Female——A much paler species than nigrifemur. Head coloured as in that
species, but the palpi are brownish apically, and the hairs on middle of cheeks are
yellow, not black. Thoracic vittae anteriorly, humeral spot, and lower portion of
sternopleural mark reddish. Apices of abdominal tergites broadly pale yellow. All
femora and the hind tibiae with a black partial annulus at middle, most distinct on
dorsal surface. Wings clear. Halteres yellow.

Structurally the same as in nigrifemur, the hind femora about three times as
thick as the middle pair, but the cheeks are not so high, and the second costal
section is distinctly longer than the third.

Length, 3-5 mm.

Type, National Park, Tasmania, 17th Dec., 1922 (A. L. Tonnoir).

CHLOROMERUS GRACILIS, N. Sp.

Male.—Head clay-yellow, the triangle glossy brownish yellow, ocellar spot,
central half of upper occiput, aristae, and third antennal segment, except its base
on inner surface, black; palpi yellow; clypeus brownish yellow. Thorax glossy
testaceous yellow, darker on dorsum, mesonotum with the usual five glossy black
vittae, and a dark spot on each humerus; pleura with only the spot on lower
portion of mesopleura deep black, the one on sternopleura blackened above, those
on pteropleura, hypopleura, and below prothoracic spiracle, brownish; metanotum
black. Abdomen clay-yellow, more or less infuscated above, and with a black spot
on each side of second tergite. Legs honey-yellow. Wings hyaline. Halteres white.

Frons fully one-third of the head width; triangle filling vertex, its sides
straight, apex at anterior margin, surface polished, each lateral margin with a
series of fine hairs; inner vertical bristles much weaker than the outer pair; third
antennal segment suborbicular; arista microscopically pubescent; cheek not over
half as high as width of third antennal segment. Thoracic dorsum with quite
coarse piliferous punctures; scutellum convex, rounded in outline, the two apical
bristles quite long. Legs more slender than in the other species of the genus, the
hind femora not over twice as thick as mid pair, and not thicker at setulose part
than proximad of it; hind tibiae not noticeably curved. First costal division

BY J. R. MALLOCH. , 433

(the one beyond humeral vein) not two-thirds as long as second and a little
shorter than third; penultimate section of third vein about half as long as
penultimate section of fourth, the latter shorter than ultimate section of fifth.
Length, 2.25 mm.
Type, Sydney, N.S.W., 29th Jan., 1925; paratype, Lindfield, N.S.W., 26th Febr.,
1922 (A. J. Nicholson).

CHLOROMERUS TRIMACULATA, 0. SDP.

Brighter yellow than gracilis, frons yellow, with the exception of a black
triangular mark on ocellar region; antennae orange-yellow; two blackish marks
on the clypeus. Thoracic dorsum with the usual five glossy black vittae, each
humerus with a black spot; pleura with only three spots, which are entirely glossy
black, these situated on the lower part of mesopleura, on sternopleura, and hypo-
pleura, metanotum broadly black in centre. Abdomen with a large black spot on
each side of second tergite, and bases of other tergites blackened. Legs yellow,
fore tarsi slightly browned. Wings hyaline. MHalteres yellow.

Frontal triangle glossy, occupying almost the entire vertex, and extending to
anterior margin of frons, its sides with some fine hairs, but not sulcate; cheek
over half as high as width of third antennal segment. Thoracic dorsum with
numerous piliferous punctures. Fore tarsi not dilated; hind femora about three
times as thick as mid pair. First costal division fully two-thirds as long as
second, and about equal to third; penultimate section of third vein a little shorter
than penultimate section of fourth, the latter shorter than ultimate section of
fifth. !

Length, 2-75 mm.

Type, Barrington Tops, N.S.W., 25th Jan., 1922 (A. J. Nicholson).

CHLOROMERUS PURUS Becker.
I have distinguished in the foregoing key three forms of this species to which
I have given names. It is possible that these may be entitled to specific rank, but
my material is too scanty to permit of a full investigation of this point.

CHLOROMERUS PURUS, form PURUS Becker.

In this form the mesonotal vittae are either entirely black or there is a small
portion at or near the anterior extremity of the central vitta reddish. In addition
to this character the hind tibia has usually a dark dorsal mark near middle, the
humeri are unspotted, and the frontal triangle always has a black marginal line
on each side. The first costal division is distinctly longer than third.

Locality, Kosciusko, N.S.W., 27th Nov., 1922, and 7th Dec, 1922 (A. J.
Nicholson). ;

CHLOROMERUS PURUS, form MACULIFERA, novum.

This form is distinguished from the typical form in having a blackish mark on
underside of each femur at middle, and a black spot on each humerus. The hind
tibia is dark marked as in purus. Penultimate section of fourth vein longer
than ultimate section of fifth.

Type, Lake Margaret, Tasmania, 3rd Feb., 1923 (A. L. Tonnoir).

CHLOROMERUS PURUS, form VARIANS, novum. :
This form has the dorsal vittae of the thorax largely, sometimes entirely,
reddish, usually the black markings confined to posterior and lateral portions.

434 NOTES ON AUSTRALIAN DIPTERA, Xlil,

Type, and one paratype, Kosciusko, N.S.W., 7th Dec., 1922, two paratypes.
Perth, W.A., 15th Nov., 1924 (Nicholson) ; two paratypes, Cradle Valley, Tasmania.
12th Jan., 1923 (A. L. Tonnoir).

In all probability many more species of this genus will be discovered by
careful collecting and being an indigenous genus it will be of interest to find
out what are its larval habits.

Genus DipLtotoxa Loew.

This genus is similar to Chlorops Meigen, but has the cross-veins of the
wings much closer together. In the only species of the genus amongst the
material now before me from this region, the third antennal segment is short,
and rounded at apex, but some of the American species have the third antennal
segment much longer than wide. This latter segregate is the typical one,
containing as it does the genotype, versicolor Loew, occurring in North America.

DIPLOTOXA TASMANIENSIS, n. sp. (Text-fig. 9.)

Male and female.—Clay-yellow. Frontal triangle wholly glossy, the colon
brownish yellow, ocellar spot black; third antennal segment slightly darkened
above; aristae, centre of occiput above, and the clypeus, black. Thoracic dorsum
with three broad black or brownish vittae which are slightly grey dusted, the
median one not extending to posterior margin of mesonotum, and two. narrow
vittae behind suture laterad of these; metanotum broadly black in centre; scutellum
paler in centre than on sides; pleura with five black marks. Abdomen more or
less browned at bases of tergites. Legs without noticeable dark markings. Wings
hyaline. MHalteres yellow. |

Frontal triangle extending narrowly to anterior margin of frons; third
antennal segment not longer than wide; arista thinner than in microcera Loew,
and a little longer, the pubescence very short. Thorax normal. Legs slender.
Wing as in Text-fig. 9.

Length, 1:75 mm.

Type, male, and allotype, Eaglehawk Neck, Tasmania, 14th Nov., 1922 (A. L.
Tonnoir).

This species is very similar to microcera Loew, a North American species
that occurs commonly in marshy spots in the central states, but in the latter
the frontal triangle is not polished and it does not extend to anterior margin of
frons. Nothing is known of the larval habits of microcera.

I had intended to publish in this part of the series a key to the genera of
Chloropidae but lack of time to finish the work on the Australian material causes
me to defer this until a subsequent part, though it is possible now to present a key
to the genera of the subfamily Botanobiinae, with keys to certain genera, herein.

Subfamily BoraNoBIIN AE.

The members of this subfamily are distinguished from those of Chloropinae
by the continuation of the costal vein to apex of fourth vein, and the invariable
presence of a sensory area on the dorsal surface of the hind tibia.

In presenting the following key to genera I realize that there is a very
great probability that it is far from complete as the material in my hands is not
sufficient to warrant one forming the opinion that all Australian genera are
represented in it, and the area covered by the collections is comparatively small.
Later should there be material forthcoming to permit a fuller treatment I will
publish a more complete synopsis.

bo

(JS)

-1

10.

Hila

13.

BY J. R. MALLOCH. 435

Key to the Geneva.

Fifth wing vein absolutely straight along discal cell .......... Biocerencia chro ao eenCoe 2
Fifth wing vein with a more or less conspicuous flexure or bend at about middle of
AiSCalMCe lle wart! cue netenelscusle/rorreu ave liavromacaise te senawene suahial casi/s folleclsasital <6 Semon aielicd san oweitey serchekts 4
Second wing vein very short, ending in costa close to apex of first vein, the second
costal section not half as long as third ................ Siphunculina Rondani.
Second wing vein not exceptionally short, ending in costa far from apex of first
vein, the third section of costa not longer than second ...................-.- 3

Seutellum about twice as long as its basal width, gradually tapered from base to
apex, where it is about one-fourth as wide as at base, with a broad shallow
central dorsal sulcus; mesonotum trisulecate; hind tibial spur quite large. ......

Bod eo tr A eu GuG IO CUO IO! DHS URIS, 0 otal Hebe ben ONE 4a ac grE Sua Ue rei aOR Resse Euhippelates Malloch.
Scutellum not longer than its basal width, rounded in outline, and not sulcate;
thorax not suleate on dorsum; hind tibial spur microscopic .... Platyina, n. g.

Humeri each with two bristles, the inner one curved towards middle of dorsum,
the outer one curved backward; dorsocentral bristles on thorax -usually quite
long, and more than two postsutural pairs present .....................-. 5

Humeri with no inwardly directed bristle ................. cece cet eee sce vces 6

Arista normal in form in both sexes; hind tibia always with a more or less evident
apical spur on anterior side, sometimes very small; scutellum with two or more
fine discal bristles or hairs in addition to the marginal bristles ............
ones cla Ars ost sg Uonteslcd ecru feum lel valiant aug Seay aue Noe eb oon emer eh aera alee -...... Parahippelates Becker.

Arista normal in form in the female, with the second segment much elongated and
almost as long as third in male, geniculated at the division between these
segments; hind tibia without an evident apical spur; scutellum without discal
hairs, only the marginal bristles present .............. Ephydroscinis Malloch.

Hind tibia with a more or less noticeable black spur at apex on anterior side P
EN Reyes rahe chaise oRic oak Sere Wedted(otin eagslbe Wapeuie cure We ulctc aise asic e aictee oie ts le tsoruehiays oA Side en oust owe wane to eile Hippelates Loew.

Hind tibia without any apical spur on anterior side .....................20000- 7
Mesopleura quite densely haired on its upper posterior portion; scutellum rounded
TOUT Ege year Nene ASeL Mela ecKs A Ree ry SoyNR er udeae ‘Batrachomyia Skuse.
Mesopleura without distinct hairs, usually pollinose or microscopically tomentose
ONMUPVDEHMPOSCETIOLMYOLLIONI A enecsie lense, eheleconenayeiclicte eect ce ae Ceeane Meee ee rciene cudinge ole 8

Frons flattened and precipitous from above middle, the posterior third horizontal,
triangle not extending beyond the horizontal part, antennae situated well below

middle of eye in profile; notopleural bristles 1 +1 ........ Benjaminella Malloch.
Frons horizontal, or sloping downward, on entire length, antennae usually inserted
at or above middle of eye; notopleurals 1 + 2 or 14 3......... Scr onie cor et aiie rene 9

Scutellum longer than its basal width, flattened above, tapered to apex, and with
some setigerouSs warts on margin apically; mesopleura tomentose on upper
MOStEFION POLCLOMMes s.5 Moeals ceateiss Meee nates chsieieee o oleae RE EM Thyridula Becker.

Scutellum not longer than its basal width, and without ‘marginal setigerous warts 10

Face sharply carinate in centre, the anterior outline of mouth-opening seen from
below in the form of a wide V; penultimate section of fourth vein fully six

times as long as penultimate section of third ............ Deltastoma Malloch.
Face with or without a central carina, but in all cases with the outline of mouth-
opening forming a transverse or arcuate line anteriorly .................. 11

Frontal triangle not defined beyond immediate range of ocelli; scutellum flattened
above, slightly tapered to apex, where it is narrowly transverse and. furnished
with two short stiff bristles, the disc with short spinules; face deeply concave

PMR ence Nes ets hemaried preted tire citer eiletavis te) eitel wuatanse ers Sibpodooudocagooscoscosun GGinagng Mrliodn
Frontal triangle usually well defined, and always extending much beyond range of
ocelli or the scutellum and face are not as above .................000ceee 12
Thoracic dorsum with three punctate linear furrows or sulci ...... Tricimba Lioy.
Thoracic dorsum without well defined sulci or furrows ........0.....000ee0e eee 3

Arista quite densely short haired, the longest hairs longer than its basal. diameter
Prd Ciccolo kaon shoe eae ChloboonO donde Qe Addu DOdeDOOOdSCODO OK OOOH OO eS. ~MEGOKRNAR IUCN,
Arista bare or pubescent, the longest hairs not longer than its basal diameter
POMS CHeM NNSA Mem WER ie raliay el site aires ietseviel autawes aivell cite Volsshawene hei erie a ero) enahsmererenee at aareh ane *Botanobia Lioy.

* The line of demarcation between these two genera is not very sharply drawn.

436 NOTES ON AUSTRALIAN DIPTERA, Xiii,

Genus SIPHUNCULINA Rondani.

I have described an Australian species of this genus, breviseta Malloch, from
New South Wales. The genotype, which is possibly cosmopolitan, ought to occur
also.

Genus EUHIPPELATES Malloch.
The genotype, pallidiseta Malloch, was described from New South Wales.

Genus PLATYINA novum.

A peculiar genus, with no flexure of the fifth vein on its penultimate section.
The cross-veins are quite noticeably approximated (Text-fig. 10), the frons and
thorax are more flattened than usual, and the face has a central carina which
widens below and above, leaving two lateral rounded cavities in which lie the
divergent antennae (Text-fig. 11). Genotype, P. nebulifera, n. sp.

Text-fig. 10. Platyina nebulifera, wing.
Text-fig. 11. Platyina nebulifera, head from front.

PLATYINA NEBULIFERA, Nn. Sp. (Text-figs. 10 and 11.)

Female.—Head testaceous, occiput and upper two-thirds of frons black, the
whole greyish dusted; third antennal segment brownish above; aristae fuscous;
palpi testaceous. Thorax black, upper parts of pleura and entire dorsum with
greyish dusting, but quite noticeably shining, the short dorsal hairs yellow.
Abdomen above concolorous with mesonotum. Legs testaceous, all coxae and
femora almost entirely blackened, and usually at least the mid and hind tibiae
browned at middle. Wings whitish hyaline, clouded centrally as in Text-fig. 10.
Halteres white.

Frons nearly half of the head width, with short regular hairs, slightly
projecting in profile, the vertical and postvertical bristles very short, ocellar
bristles microscopic; third antennal segment not as long as wide; arista pubescent;
cheek fully as high as width of third antennal segment; eyes haired, longer than
high; palpi quite large. Thoracic dorsum smooth, with quite dense decumbent
microscopic hairs, the humeral bristle as long as the notopleurals, the latter 1 + 2;
scutellum wider than long, narrowly subtransverse at apex, the disc with micro-
scopic hairs, the apex with two small incurved setulae. Legs normal. Wing
venation as in Text-fig. 10.

Length, 1:75 mm.

Type and one paratype, Eaglehawk Neck, Tasmania, 14th Nov., 1922 (A. L.
Tonnoir).

I am not absolutely certain of the sex of these specimens but think that
they are females.

BY J. R. MALLOCH. 437

Genus PARAHIPPELATES Becker.
I have in preparation a new synopsis of the species of this genus, the one
previously published by me (These ProcreEpiINGs, 1923, p. 619) being incomplete
owing to the subsequent description of some species.

Genus EpHyDROSCINIS Malloch.

There is but one known species of this genus, australis Malloch.

Genus HIPPELATES Loew.

There are several undescribed species of this genus in my hands now and
below I present a key for the recognition of the species known to me up to this
time.

It would be of interest to know whether the Australian species of this genus
have the same habits as some of those occurring in America, where the adults
are very persistent in flying about persons in the open, apparently attracted by
perspiration, and they have been considered as transmittors of certain diseases
of the eye.

Key to the Australian species.

1. Third antennal segment entirely deep black; hind tibial spur brownish yellow, not
as long as the apex of the rather thick tibia, quite stout and curved, situated
close to apex; mesonotum with three glossy black vittae on a yellow ground,
the central one complete, the laterals abbreviated at both extremities, and

slighthysunterrupted@ata SULUTE Murry aleictersieliceier Scienia welciiehe eines atricornis, n. sp.
Antennae either entirely yellow or with a very slight darkening of the upper edge
of third segment; hind tibial spur deep black ............... saat stoanstie. sisi cnenshele says 2

2. Hind tibial spur much curved and distinctly longer than tibial diameter, situated
well in front of apex of tibia; thoracic dorsum with a large quadrate mark
in centre of hind margin, and a spot near each humeral angle, deep black, the

Exo IGl Golo e AVEWO oso Soop obaob oO Un ULoNs GoD oO DIMOICGOG5 ao bancrofti Malloch.
Hind tibial spur not as long as tibial diameter, slightly curved, and situated at,
or very close to, apex of tibia; thorax not marked as above ................ 3

8. Mesonotum and disc of scutellum with rather coarse setigerous punctures, the
- scutellum slightly rugose, disc of mesonotum black, the usual three black vittae
fused on their entire length leaving only the lateral margins yellowish, the
postsutural lateral vittae separated from the central black complex; scutellum
broadly blackened across base; sternopleura entirely glossy black ..........

RI GR sh 1 od, Tk a NR OO NEW AC = et RO eNO co nigridorsata, n. sp.
Mesonotum and dise of scutellum with a few weak setigerous punctures, sparsest on
scutellum; disc of mesonotum with distinct black vittae; scutellum not blackened

across base; sternopleura partly, or entirely, yellow .......................-: 4

4. Pleura absolutely without black markings, entirely yellow; thoracic dorsum marked
as Text-fig. 12, d; frontal triangle glossy black ............ atriseta (Malloch).
Pleura with at least one black mark; thoracic dorsum not marked as above .... 5

5. Sternopleura largely black in centre, two black spots on mesopleura, and one on
pteropleura; discal thoracic markings as Text-fig. 12, a@ ...... abbreviata, n. sp.
Sternopleura yellow, only one black mark on pleura .......................... 6

6. Discal thoracic markings as in Text-fig. 12, b; tibial spur moderate ..........
SOO 6005 0100 HONS DID SOTHO OOOO HHODOLADOOLOO dO dood GOO DRONA OHIO Glo. S10:
Discal thoracic markings as in Text-fig. 12, c; tibial spur minute .. fergwsoni, n. sp.

The first two species dealt with are distinguished by having the hind tibial
spur extremely small and inconspicuous, so much so in fact that I originally
described one of them as a species of Gauraz. I believe, however, that they are
properly referable to Hippelates.

438 NOTES ON AUSTRALIAN DIPTERA, Xiii,

HIPPELATES ATRISETA (Malloch). (Text-fig. 12, d.)

Gaurax atriseta Malloch, Proc. Linn. Soc. N.S.W., 1924, xlix, p. 355.

Originally described as a species of Gaurax. Readily distinguished from
any of those now included in the genus by the entirely yellow pleura and the
inconspicuously marked thoracic dorsum (Text-fig. 12, d).

Locality, Sydney, N.S.W.

Text-fig. 12. Dorsal thoracic markings of species of Hippelates. a, abbreviata ;
b. unimaculata; c, fergusoni; d, atriseta.

Text-fig. 13. Tricimba carinifacies, head from front.

Text-flg. 14. Tricimba pollinosa, head from front.

HIPPELATES FERGUSONI, nh. sp. (Text-fig. 12, c.)

This species is fulvous yellow in colour, with the frons and scutellum more
lemon-yellow, the frontal triangle with a central dark vitta, third antennal segment
with a dark mark at insertion of arista, thoracic dorsum marked as in Text-fig.
12, c, the pleura with but one black mark, on lower margin of mesopleura, the
abdomen black on dorsal surface except on first tergite, centre of second, and the
sides of fifth.

Structurally the species is very similar to atriseta, but the lateral hairs on
frons are stronger.

An additional character is that the palpi are brown instead of yellow, but
the colour may be due to some unnatural cause, and abnormal.

Length, 2-5 mm.

Type, Sydney, N.S.W., 26th Aug., 1923.

I take pleasure in dedicating this species to the late Dr. E. W. Ferguson, to
whom I was indebted for most of the material used in the compilation of this
series of papers.

HIPPELATES ATRICORNIS, N. SD.

Female.—Head yellow, with yellow hairs and brownish bristles; ocellar spot,
third antennal segment, and aristae, black. Thorax yellow, dorsum with three
glossy black vittae, of which only the central one extends the entire length, and
traces of two narrow reddish vittae laterad of these behind the suture; pleura
with a black mark on each of the following parts: mesopleura, pteropleura, sterno-
pleura, and hypopleura; scutellum yellow; metanotum broadly glossy black in
centre; hairs pale, the bristles black or brown. Abdomen yellow, with a black
suffused spot on each side of second tergite, third and fourth tergites almost
entirely black on disc, fifth with a central black spot. Legs yellow. Wings hyaline.
Halteres yellow.

A

BY J. R. MALLOCH: 439

Frons longer than wide, quite copiously fine-haired, the lateral margins with
a series of short setulae, vertical and postvertical bristles quite well developed,
ocellars short; eyes hairy, higher than long; cheek not half as high as width of
third antenal segment, the latter orbicular; arista with very short pubescence;
proboscis stout, geniculated. Thorax with quite long fine decumbent dorsal hairs,
the surface hardly punctate, and the bristles well developed; scutellum slightly
flattened above, with two long and two short apical black bristles. Legs stout,
the hairs quite long, apical spur on hind tibia stout, curved, and about as long
as diameter of tibia. Penultimate section of fourth vein about twice as long
as penultimate section of third, and about equal in length to ultimate section of
fifth.

Length, 3 mm.

Type, Cairns, N. Queensland (J. F. Illingworth).

Type in United States National Museum.

HIPPELATES NIGRIDORSATA, Nl. SD.

Female.—Head orange coloured, occiput in centre, and the ocellar region, black,
a brown streak running forward from ocellar spot to front of triangle; clypeus
black; antennae yellow; aristae black. Thorax orange-yellow, disc marked as
stated in key, humeral angles unspotted; pleura black except in front and above.
Abdomen brownish orange. Legs dirty orange-yellow, a dark spot near base of
each fore coxa, and some dark suffusions on hind femora which may be abnormal.
Wings hyaline. Halteres yellow.

Frons a little less than half the head width, triangle shining, not conspicuously
differentiated on sides, microscopically striate laterally; ocellar bristles minute,
postvertical pair much larger; orbitals distinct posteriorly; face concave, with
a V-shaped elevation above between bases of antennae; eyes hairy; cheek about
one-eighth of the eye height, not much produced anteriorly, with black hairs
below; palpi elongate, not very slender; proboscis stout; antennae normal; arista
pubescent. Mesonotal hairs short; subapical pair of bristles on scutellum much
shorter than the apical pair. Legs normal, apical spur on hind tibia not half
as long as the diameter of tibia, situated close to apex. Second section of costa
a little longer than first and about 1-75 as long as third; apical sections of veins
3 and 4 parallel.

Length, 4 mm.

Type, King River, Tasmania, 4th Febr., 1923 (A. L. Tonnoir):

HIPPELATES UNIMACULATA, TD. Sp. (Text-fig. 12, b.)

Male.—The general colour of this species is paler than that of nigridorsata,
the lateral margins of mesonotum, and especially the disc of the scutellum, being
conspicuously so, lemon yellow. The abdomen is, however, extensively blackened
on dorsum, the first visible tergite having a black spot on each side, the second
and third largely black, only yellow on anterior and posterior margins laterally,
and the fourth black except on its anterior lateral margins. Legs yellow, a small
black spot behind on base of each mid coxa, and there is a black mark on middle
of frontal triangle before ocelli. Wings clear. Halteres yellow.

Frons narrower than in nigridorsata, narrowest anteriorly, triangle glossy,
well defined, extending to anterior margin of frons, orbital setulae rather strong;
other characters as in nigridorsata. Scutellum more flattened than in that species,
apex very slightly pointed between the apical bristles, the latter longer than in

440 NOTES ON AUSTRALIAN DIPTERA, X1iii,

nigridorsata, the subapical pair more remote from apical pair and not so short.
Apical spur on hind tibia fully half as long as tibial diameter, rather stout,
close to apex. Venation similar to that of nigridorsata.

-Length, 3-5 mm.

Type, Burnie, Tasmania, 5th Oct., 1922 (A. L. Tonnoir).

HIPPELATES ABBREVIATA, . Sp. (Text-fig. 12, a.)

Male.—Similar to wnimaculata in colour, the scutellum lemon-yellow, and the
abdomen with lateral dark spot on first visible tergite and a dark apical fascia
on each of the other tergites. Distinctions as in key.

The frontal triangle extends only two-thirds of the way to front margin of
frons; the cheeks are narrower than in wnimaculata; the apical spur on hind
tibia is shorter, and the species is not so large.

Length, 2 mm.

Type, Burnie, Tasmania, 5th Oct., 1922 (A. L. Tonnoir).

Genus BatTracHomytA Skuse.

I have already described two species of this genus from Australia and having
two new species from Tasmania present herewith a key for the identification of
the four. The genotype is unknown to me, but appears to be distinct from these.

Key to the Species.

1. Thoracic dorsum rufous yellow, with four deep black vittae ................... 2
Thoracic dorsum brownish or yellowish, without distinguishable vittae ............ 3

2. Notopleural area and scutellum not much paler than remainder of thoracic dorsum,
the scutellum not blackened on basal lateral angles; humeri without a black

Spots tibiae largely: fuscousp te ace areca eliche icra Casta SES one me iaeiey oie sue major, n, sp.
Notopleural area and scutellum ivory-white, the latter blackened on basal lateral
angles; humeri with a black spot; tibiae yellow .............. sStrigipes, n. Sp.

3. Third antennal segment deep black; femora largely black-haired .. atricornis Malloch.
Third antennal segment yellow; femora entirely yellow-haired .... flavicornis Malloch.

BATRACHOMYIA MAJOR, 1. SD.

Male and female.—Head yellow, opaque, variegated with brown, the frons
sometimes brown except on anterior margin; clypeus blackish; antennae and
palpi yellow, third segment of former more or less darkened at insertion of arista;
aristae black; cephalic hairs black except on lower part of occiput. Thorax
reddish yellow, shining; mesonotum with three broad black vittae which are
fused on anterior half, and a narrow postsutural black streak which is separated
from the submedian vitta; postnotum black; four large black spots on pleura,
on lower part of mesopleura, on pteropleura, on hypopleura, and on all except the
upper margin of sternopleura. Abdomen rufous yellow, with the apices of
tergites darkened. Legs rufous yellow, the fore femora and all tibiae almost
entirely black. Wings slightly greyish. Halteres vellow. Fine hairs on thorax
mostly pale, the bristles, and many of the hairs on abdomen, black.

Frons about two-fifths of the head width, the surface with many black hairs;
post-vertical bristles long; ocellar pair shorter, both pairs incurved; antennae
narrowly separated at bases, third segment higher than long; arista rather thick
basally, microscopically pubescent; cheek about one-fifth of the eye height, rather
densely haired except on upper margins, the hairs ascending well above the mouth
margin te vibrissal angle; palpi normal; eyes densely haired. Dorsum of thorax
including that of scutellum quite densely haired, the hind margin of scutellum
with a number of setulae. Abdomen more hairy than usual, but not so densely

BY J. R. MALLOCH. 3 441

so as mesonotum. Legs rather robust, tarsi slightly broadened apically. First
section of costa subequal to second in the male, a little longer than it in female,
third section about half as long as second in- male, distinctly less than that in
female.

Length, 4-5-5-25 mm.

Type male, allotype, and four paratypes, Cradle Valley, Tasmania, 10th to
26th Jan., 1923 (A. L. Tonnoir).

BATRACHOMYIA STRIGIPES, N. SD.

Male.—A paler species than major, the head, except ocellar spot, a part of
occiput, and aristae, pale yellow; mesonotal vittae not fused anteriorly, connected
at hind extremities. Wings hyaline. Haltéres cream coloured.

Hairs not so dense as in major; third section of costa about two-thirds as
long as second, the latter subequal to first.

Length, 3-5 mm.

Type, Wilmot, Tasmania, 8th Jan., 1923 (A. L. Tonnoir).

Genus BENJAMINELLA Malloch.
There is but one species of this genus known, albifacies Malloch, which occurs
in Tasmania and New South Wales.

Genus THYRIDULA Becker.
There are three species already described from Australia, and I now add a
fourth one, which is as yet undescribed. Below I present a key for the identifica-
tion of the species, and a description of the new one.

Key to the Species.

1. Scutellum paler at apex than at base, quite coarsely rugoso-reticulate on entire
surface, its sides parallel on basal two-thirds of their extent, the apex broadly
rounded, and with four subequal bristles situated on small warts; claws of
hind tarsi not much longer and stronger than those of mid and fore tarsi
SORE EO EEOLE -O30..5..0 \0v0| ONS. GO ADE BEGG SERGE yall CHC CANC RGEC Peta CCR eRe cee Gackt rugosa Malloch.

Scutellum darker at apex than at base, or with a black central vitta, and not
coarsely rugoso-reticulate, the sides converging from base to apex, at the latter
narrowly truncate and with a pair of quite long bristles situated on two small
warts; claws of hind tarsi much stronger than, and at least twice as large as,

those of, smidwandifone™ tarsic, gers rate eS iene cos cholerae eke re tReet cy ean nee ane Mos 2

2. Scutellum with a complete, deep black, central vitta; face broadly. blackened
(CLIMB PUNY Re o'6 4 Sig 6.c do MMOLE DICMET ELST HON OTO Gr otaitte Is te oter me Nienateee Sees centralis Malloch.
Scurellumpblackwonsaprealehalfsorslessmapeecaeciicie cai ieicaiicne he nieiiineimeincle 3

Face and hind tarsi entirely yellow; frons yellow, ocellar spot black ..............
CeO ASRD A lode che Dio OnCeEnG ra Rocce ete ulokeinct ii ena Rin BPR aco a ri area ee Mr dl te atroapicata Malloch.
Face blackened in centre; fourth and fifth segments of hind tarsi black, the other
segments yellow; frons dark brown, yellow only on anterior and _ posterior
meen, WS Ceallkye Hoos WEOK pooadccbadonuonoocouducUacKD brunneifrons, n. sp.

(Se)

THYRIDULA (EUTHYRIDULA) RUGOSA Malloch.
I erect for this species the new subgenus Euthyridula, based upon the
characters cited in the key.

THYRIDULA CENTRALIS Malloch.
This species is known only from the type specimen from: Sydney, N. S. Wales.

THYRIDULA ATROAPICATA Malloch.
This species is known only from the type specimen from Bowral, N. S. Wales.

442 NOTES ON AUSTRALIAN DIPTERA, Xiili,

THYRIDULA BRUNNEIFRONS, N. SD.

Male.—Head brownish testaceous, centre of face blackish, frons dark brown,
yellowish only on anterior and vertical margins, the ocellar spot glossy black;
antennae testaceous yellow, third segment brownish except at base; aristae fuscous;
palpi testaceous yellow. Thorax brownish testaceous, dorsum with three black
vittae, a complete one in centre, and one on each side of it which is not carried
to anterior nor posterior margin and is not interrupted at suture, and laterad
of these behind suture a black streak; humeri unspotted; sternopleura mostly
black, a black spot on lower part of mesopleura, one on pteropleura, one behind
prothoracie spiracle, and one on hypopleura; scutellum black at about the apical
half: metanotum glossy black. Abdomen testaceous yellow, with a dark mark
on each side of at least the second tergite. Legs testaceous yellow, mid and hind
femora and tibiae more or less darkened on middle; fourth and fifth segments
of hind tarsi black. Wings hyaline. Halteres pale yellow.

Frons a little longer than its anterior width, quite densely haired, the orbital
setulae hardly differentiated, triangle not extending to middle of frons; third
antennal segment quite large, orbicular; arista distinctly pubescent; face slightly
carinate above, distinctly produced below; eyes densely haired; palpi long;
proboscis long, geniculated, and slender, the apical section as long as head; cheek
almost linear. Thorax as in centralis. Legs stout, hind tarsal claws fully twice
as long as those of mid and fore pairs.

Length, 2 mm.

Type, Strahan, Tasmania, 6th Febr., 1923 (A. L. Tonnoir).

Genus DELTASTOMA Malloch.
Only one species of this genus is known, unipunctata Malloch; Sydney, N.S.W.

Genus Cavicers Malloch.
Only one species of this genus is known, flavipes Malloch; Sydney, N.S.W.

Genus TrRICIMBA Lioy.

Before me are four species of this genus which appear to be undescribed.
All of them are very different from the genotype in structure, two having a very
conspicuous flat carina on upper half of the face separating the antennae quite
widely, and the frons quite noticeably protruded in front, projecting well over
antennae, much as in the genus Eurina Meigen. One of the other species is much
less robust than the genotype, without a facial carina, and with larger wings.

One might be justified in erecting a new subgenus for the reception of the two
first mentioned species but for the fact that an Australian species, carinata
Malloch, to a considerable extent bridges the gap between these and the genotype.
In the Australian species the facial carina is present, but not so wide, and the
frons is less protruded in front. The species now known to me are included in
the key below.

Key to the Species.

1. Face with a quite broad conspicuous central vertical carina on upper half ........ 2
Nace not-atvalleevidently: Carin ata ay. ype ons caccl shone cists te ioe Mens oils wre colle ekeuchepeneeedoles sateen ty ue
Small species, not over 2 mm. in length; frons not projecting beyond anterior margin
of third antennal segment in profile; facial carina rather narrow, not widened
out at lower extremity, and extending well below middle of face ............
PAE mR BLE Ch OO OuCLCAD CRORE REREACHCADERO CLD ECIC IY EAEIOLE CT CU BOLING ote hcubach ch acolo carinata Malloch.

BY J. R. MALLOCH. 443

Larger species, not less than 3 mm. in length; frons projecting well beyond anterior
margin of third antennal segment in profile; facial carina broad, distinctly
widened out at lower extremity, and not extending below middle of face .... 3

Sternopleura grey dusted on upper and lower margins, the upper margin testaceous,
THERLECNAINGCEMDIAC Kea aivarsicveroricrs elclercucicisrer stn otal aah siclels vaicuesensrane carinifacies, N. sp.

ie)

Sternopleura entirely grey dusted, and all black .................... pollinosa, n. sp.

4. Scutellum quadrate, flat on entire dorsal surface, and with a prominent setigerous
wart near each posterior lateral angle; small species, not over 1:5 mm. in

IEG A, WAG: WENA TROL soacesadnobantoscusodaodcuonbDonEdoaODOGO UCD OcOS 5

Secutellum longer than wide, tapered to apex, the sides sloping basally, and the margin
without setigerous warts; species averaging fully two mm. in length, and of
Piss re Adal NeiSin Go Solos cua due Ore La DO ba 6 Cola Dore a drotorotoer or Orecore convexra, Nn. sp.

Mesonotum with but one evident series of microscopic pale hairs between the central
and lateral sulci; frontal triangle with evident grey dusting, but distinctly
shining, ending a little in front of middle of frons, and with no wedge-shaped
mark in front of it, the anterior margin of frons orange-yellow; legs testaceous
yellow; penultimate sections of third and fourth veins subequal in length
NS eC a OO OEE SIRO CIOL IO OS CIOS ISOC IONE OF CRE Le Did onGarel CPOE ERENCre scutellata Malloch.

Mesonotum with about four series of microscopic dark hairs between the central
and each lateral sulcus; frontal triangle densely grey dusted, and not shining,
ending at middle of frons, but with a wedge-shaped pale-dusted mark in front
of- it which gives it the appearance of being continued to anterior margin of
frons, the latter hardly paler than remainder of surface; mid and hind femora,
and hind tibiae, with a brownish median mark; penultimate section of third vein
about half as long as penultimate section of fourth .............. similata, n. sp.

ol

TRICIMBA CARINIFACIES, n. sp. (Text-fig. 13.)

Female.—Head yellowish testaceous, becoming fuscous on upper part of frons
and occiput, third antennal segment dark above; clypeus fuscous. Thorax
brownish testaceous, dorsum fuscous, with grey dusting, but distinctly shining
on most of the surface, lower part of sternopleura and postnotum black, the
latter shining. Legs testaceous yellow, femora mostly, and tibiae in middle,
brown or fuscous. Wings hyaline. Halteres yellow. All hairs and bristles luteous.

Frons projecting beyond eye fully as far as height of cheek, the latter over
one-third as high as eye; arista subnude; front view of head as in Text-fig. 13.
Mesonotal hairs stubby, lateral sulci broadened posteriorly; scutellum flattened
above, with rather dense stubby bristles or hairs on disc, and with two approxim-
ated bristles at apex. Legs normal. First costal section about four-fifths as long
as second, third two-thirds as long as it.

Length, 3-5 mm.

Type and one paratype, Burnie, Tasmania, 25th Oct., 1922 (A. L. Tonnoir).

TRICIMBA POLLINOSA, n. sp. (Text-fig. 14.)

This species is very similar to carinifacies, but is smaller, and darker in colour,
the frons is darker, being sometimes entirely fuscous, and the sternopleura is
entirely grey dusted. In addition to the characters listed in the key the face is
narrower, being at lower extremity of the parafacials not wider than the height
of cheek, while in carinifacies it is distinctly wider, and the carina in this species
does not extend to middle of face (Text-fig. 14), while in carinifacies it does
extend to, or even very slightly below, the middle.

Length, 3 mm. :

Type male, Woy Woy, N.S.W., 2nd Sept., 1923; one paratype, Lindfield,
N.S.W., 19th Nov., 1922 (A. J. Nicholson); one paratype, Bridgetown, W.A., 29th
Aug., 1926 (E. W. Ferguson).

444 : NOTES ON AUSTRALIAN DIPTERA, Xiil,

The Western Australian specimen is slightly different from the others, but
not so distinctly so as to justify me in accepting the distinctions as of specific
value.

TRICIMBA CONVEXA, ND. SDP.

Female.—Differs from carinifacies in having the frons darker, the thorax
and abdomen uniformly fuscous, shining, with greyish dust except on lower half
of pleura, and the legs uniformly yellowish testaceous.

The frons is not half of the head width, the cheeks are almost linear, the
mesonotum is more convex, with finer hairs, and the lateral sulci are not widened
posteriorly, the legs are more slender, and the wings broader and longer. }

Length, 2-2-5 mm. ye ;

Type and one paratype, King River, Tasmania, 4th Febr., 1923; one paratype,
National Park, Tasmania, 6th Dec., 1922; two paratypes, Adventure Bay, Tasmania,
Dec., 1922 (A. L. Tonnoir).

The most slender species of the genus known to me.

TRICIMBA SIMILATA, nN. Sp.

This species is so very similar to scutellata Malloch, which I recently described,
that it appears unnecessary to describe it further than has already been done in the
foregoing key. I have included all the essential differentiating characters therein,
and a comparison of the specimens one may have, with this key and the description
of scutellata ought to be sufficient to permit an accurate identification.

Length, 1-5 mm.

Type, Sydney, N.S.W., 9th Nov., 1924.

TRICIMBA CARINATA Malloch.
Proc. Linn. Soc. N.S.W., 1924, p. 356.
I have seen only the type female from Como, N.S.W., December, 1923.

TRICIMBA SCUTELLATA Malloch.
Proc. Linn. Soc. N.S.W., 1925, p. 337.
I have seen only the type male from Sydney, N.S.W., 9th Aug., 1924.

Genus GAauRAx Loew.

As already stated by me this genus is very poorly differentiated from
Botanobia Lioy, only the more distinctly haired arista serving to separate it from
that genus. When I come to publishing the key to the species of Botanobia I may
decide to include in it all the species of both so that no errors may occur in
identifications. I have before me a very pretty undescribed species which appears
to be referable here.

GAURAX APICIPUNCTATA, Nl. SDP.

Male and female.—Head orange-yellow, occiput, frontal triangle except extreme
apex, and upper extremities of frontal orbits, glossy black; third antennal segment
lemon yellow; aristae and their hairs black; palpi yellow. Thorax glossy black,
humeri and lateral margins of mesonotum and upper margin of pleura, fulvous
yellow; scutellum black. Abdomen rather variable in colour, ranging from
brownish to black. Legs yellow, all femora and tibiae blackened medially. Wings
hyaline, with a fuscous spot at apex from a little before tip of third to just
beyond tip of fourth vein which has usually a pale central spot. Halteres yellow.

BY J. R. MALLOCH. 445

Frons over half of the head width, with quite long black hairs; postvertical
and ocellar bristles erect and convergent; triangle wide, extending fully two-thirds
of the length of frons; third antennal segment higher than wide; arista thickened
at base, very densely short haired and appearing thickened on entire length, the
longest hairs about as long as the thick second segment; eyes hairy, a little higher
than long; cheek almost linear. Thoracic dorsum not punctate, the hairs quite
dense, long, and fine; scutellum convex, rounded in outline, with two long and
two short fine bristles. Legs stout. Second costal section fully 1:5 as long as
first, the latter a little longer than third; penultimate section of third vein over
half as long as penultimate section of fourth.

Length, 2-2-5 mm.

Type female, allotype, and one female paratype, Sydney, N.S.W.

This is the only species known to me from Australia in which the wings are
marked with a fuscous apical spot.

Genus BorTanopia Lioy.

I have been compiling a key to the species of this genus but am not yet ready
to publish it as with every consignment of specimens of small Diptera received
there are new species of this genus. Isolated descriptions are not of very much
value in many genera and this is one in which only synoptic treatment will
provide characters by means of which the species may be identified.

There appear to be about 30 species of the genus now before me from
Australia and Tasmania.

Family Asteiidae.

This family contains but three genera and may ultimately be merged with the
Drosophilidae, as it has been heretofore by many workers. Meanwhile I accept
it in the status given to it by recent workers. I have already described a species
of the genus Leiomyza from Australia and now add a species of a second genus.
Below I present a key to the three described genera. The third genus may occur
also in Australia.

Key to the Genera.

1. Outer cross-vein of the wing lacking; second wing vein ending in apex of first vein,
or in costa very close to apex of that vein; arista with quite long widely
separated hairs above and below ....................2+-eeeeees Asteia Meigen.

Outer cross-vein of wing present; arista bare or pubescent .................... 2

2. Thorax with two pairs of well developed postsutural dorsocentral bristles; second
wing vein ending in costa very close to apex of first vein; arista bare or
DUDESCET tere aoe Weer ied cay ausrinues Bionatorer oun ens nae ei ouepin ed eaverata Miner estates ons Sigaloessa Loew.

Thorax with but one pair of well developed postsutural dorsocentral bristles; second
wing vein ending in costa about midway between apices of first and third veins;
AUTISCAMDALE OL AlIMOSt, (SO. crs iicis: wileisei erie das lene eulessipesy airaeyeyeusyeneyevtets Leiomyza Macquart.

SIGALOESSA MELBOURNENSIS, n. sp. (Text-fig. 15.)

Female.—Head testaceous, a broad dark mark on each side of upper occiput,
frons reddish, face whitish, its lower rim and clypeus brown; antennae testaceous.
Thorax brownish black, shining, lightly greyish dusted on dorsum, the lateral
mesonotal margins yellowish; pleura yellowish, with two darker vittae; scutellum
yellow. Abdomen fuscous on dorsum, yellow on membranous part. Legs
testaceous yellow. Wings hyaline. Knobs of halteres dark brown above, pale
below.

446 NOTES ON AUSTRALIAN DIPTERA, Xiii.

Frons about one-third of the head width, parallel-sided, almost flat, and nearly
1-5 as long as wide, with microscopic stiff black hairs; postvertical bristles micro-
scopic, divergent as in the other genera in which they are present; both pairs of
vertical bristles rather short, but longer than the single pair of orbitals;
antennae normal; arista with very short pubescence. Thorax with a series of

os awe
> a
TNS te meen once =

15
Text-fig. 15. Sigaloessa melbournensis, wing.

short hairs on almost the entire length of the two dorsocentral lines and between
these in front of suture an irregular series of similar acrostichal hairs; posterior
pair of intra-alar setulae present; scutellum with two long apical and two short
basal bristles. Legs slender. Wing as in Text-fig. 15.

Length, 2:25 mm.

Type, Melbourne, Victoria, found in jar with Monophlebus.

Except for the dark colour in general, and especially that of mesonotum, this
species is very similar in all respects to flaveola Coquillett, a North American
species, but I consider it as distinct pending the receipt of more material from
Australia.

NOTES ON AUSTRALIAN AND EXOTIC SARCOPHAGID FLIES.

By G. H. Harpy, Walter and Eliza Hall Fellow in Economic
Biology, Queensland University, Brisbane.

(Eleven Text-figures. )
[Read 26th October, 1927.]

Taxonomic studies of Sarcophagidae, undertaken since the publication in
1923 of “A Revision of the Australian Diptera belonging to the genus Sarcophaga”,
by Professor T. Harvey Johnston and myself, were pursued with the view of
ascertaining the relationship between Australian species and those examined or
that had been described from other parts of the world. The accumulation of odd
notes so far compiled will prove, I believe, of considerable interest and perhaps
help towards establishing a satisfactory method of arranging the unwieldly genus
Sarcophaga into groups of naturally allied species.

The idea of grouping Sarcophagids on the genital characters of the male,
which has been initiated by various authors, is extended and developed in the
present paper. Already types like S. twberosa have had placed around them names
ranking as varieties or species according to the interpretations given by the
authors concerned, but unfortunately drawings of the gross appearance of the
male genitalia are usually relied upon in order to show the alliances whilst no
attempt has been made to define the associations. The genital complex of the
male is a structure worthy of detailed study and until it has been given adequate
treatment relationships between species must remain obscure.

Although highly specialized structures such as the bifid anterior clasper of
the antilope-group and the spiny anterior appendage of the peregrina-group would
be enough to suggest the alliance, reliance is not placed on such characters alone,
additional characters are taken into consideration and it is upon such combinations
that relationships are best assessed. The counterpart of any one character may
readily be found in the composition of the genitalia of species belonging to quite
different groups as instanced by the misera-group, that has one or more of its
characters found in S. securifer, aurifrons and omega. Owing to necessity for
caution in these matters only three groups are dealt with in full. Biological data
are given for one species.

For materia] supplied and used for the purpose of this paper, I am indebted to
Dr. H. H. Karny for two species bred by him in Java, the types of which by request,
are returned for preservation in the Museum there; also to Dr. R. J. Tillyard for
three specimens of the only Sarcophaga described from New Zealand, one of which
was returned to him for preservation in the Cawthron Institute; to Mr. R. R.
Parker for a collection of Sarcophagids, mostly American but including some
Indo-Australian forms; and to Mr. R. Senior-White for Oriental material in which
I was specially interested.

Method of preparing and mounting genitalia.—The published methods, whereby
the genitalia are prepared for examination (given by Parker, Senior-White, and

Johnston and Hardy), differ fundamentally and for the purpose of detecting finer
1H

448 AUSTRALIAN AND EXOTIC SARCOPHAGID FLIES,

points in the structure of genitalia in the first two cases mentioned, I find that
given by Parker is superior to that given by Senior-White. For examination of
genitalia in situ, I use a. system of reflecting sunlight on the parts examined and
any distortion or contortion is thus detected under the Greenough stereoscopic
binocular microscope, moreover the finer structural details, so often overlooked,
become clearly defined under this method. The genitalia may be removed and
mounted without distortion, rendering all external details clear on a slide; after
removal, passing through absolute alcohol, clearing in xylol or clove oil (boiling
in potash is not permissible in this case), the genitalia can be mounted whole or
in sections in Canada balsam in such a manner as not to flatten out the parts.
The parts of the penis are then easily viewed through transmitted as well as
reflected light in the position they would occupy relative to each other just as if
left set in the normal way. This method of mounting has the advantage that
higher powers of the microscope can be brought to bear upon details too small
to'be seen plainly under the binoculars, if left in situ; the densely spined nature
of the anterior appendage of NS. peregrina being a case in point.

S. antilope group.

The six species, referred to this group, agree in having the anterior clasper
bifid and with 2 small knob placed anteriorly to it. The forceps are long, strong
and angulated. The second segment of the penis lacks the lateral process, but the
shape differs in other respects. Those species that have the anterior clasper
bifid for only half the length, have also the apical process tridentate, whilst the
two species that have this clasper short and bifid to the base do not have the
tridentate character of the penis. As far as yet known the group consists of:

antilope Bottcher, 1913, Formosa.
*alpha Johnston and Tiegs, 1921, Australia.
*beta Johnston and Tiegs, 1921, Australia.
*zeta Johnston and Tiegs, 1921, Australia.
*howensis Johnston and Hardy, 1923, Lord Howe Island.
*antilopoides, n. sp., Java.
The asterisk refers to species examined.

SARCOPHAGA ANTILOPOIDES, nh. Sp. (Text-fig. 1.)

Status.—There are eleven names previously attributed to Sarcophagids
recorded from Java, and Dr. H. H. Karny has succeeded in breeding only two
species. As pointed out below, 8S. fuscicauda has been misidentified and is
described here under the name S. karnyi. 8S. taenionata, according to Senior-
White, may possibly be SNS. flavipalpis, one of the misera-group. From the same
source it is suggested that S. javana may be VS. aurifrons Doleschall (i.e. doleschali
J. and T.). Other names are lineatocollis Macquart, flavinervis Senior-White,
knabi Parker, albiceps Macquart, orientalis Parker and duxz Thomson. I can see
no evidence to suppose that antilopoides has been referred to under any of these
names, and all cf them, with the exception. perhaps, of the unknown lineatocollis
may be excluded from this consideration.

When this species was first received I considered it to be 8. antilope Bottcher,
originally described from Formosa, but later, when comparing the genitalia with
the drawing of S. antilope from India by Senior-White, I became doubtful about
the determination. Under the circumstances it seems advisable to give a new
name to the species here described which cannot be considered identical to the
Indian species and there is sufficient published evidence to conclude that the

BY G. H. HARDY. 449

Formosan species is also not the same. According to Senior-White’s drawing
there is an elongate lobe, hooklike in shape, and this appears to be indicated in the
drawing by Bottcher, but hidden between the anterior process and the sheath of
the penis. This shape is in no way indicated on any of the other forms. There
can be no doubt that the Indian and Formosan forms belong to this group but
their specific identity requires further consideration.

Description.—Male: Head, outer vertical bristles much smaller than the
postoculars; thirteen frontals, two rows of eight facials: about nine orals; one

Text-fig. 1. Male genitalia of Sarcophaga antilopoides, n. sp.—second
segment of the penis only. E, lateral view; I’, ventral view.

Text-fig. 2. Outline at the apex of the puparium of S. misera, seen
laterally.

Text-fig. 3. The same of S. kohla. :

Text-fig. 4. The last abdominal sternite on S. misera—half of the apex
only.

Text-fig. 5. Apex of the lateral process, l.p., the lobe, Jl, and the anterior
appendage, a.a.. of S. misera, seen laterally.

Text-fig. 6. The same of S. ceylonensis.

row of postoculars. Thorax, two intra-alars, the presutural and scutellar bristles
well developed. Abdomen, on the first segment one row of only a few discal
lateral bristles and one, sometimes two, submarginals, and on the third one median
pair and two or three lateral submarginals; on the fourth segment five or six
submarginals alternating with wel! developed marginals.

Genitalia: Similar to that of S. beta, but reddish, the forceps and claspers
being identical. The anterior appendage is simple but there are three minute
spine-like processes along the upper anterior edge of each half; the apical process
is rather slender towards the apex which is tridentate and has a pair of lateral
projections that are similar in position, but not shape, to those on S. howensis;
the filaments prctrude.

Legs: The chaetotaxy conforms to the general type except in the two ventral
rows on the posterior femora which are very much reduced; long hairs occur on
all femora and on posterior tibiae.

450 AUSTRALIAN AND EXOTIC SARCOPHAGID FLIES,

Female: Tie frontal bristles may be as few as eight or nine and the first
abdominal segment usually has two rows of discal lateral bristles and the ventral
bristles on the posterior femora are well developed. In other respects the
characters conform to those of the male, excluding the sexual characters.

Habitat—Java: Buitenzorg, October, 1923, 26 males, 24 females (H. H. Karny),
all bred.

SARCOPHAGA TRYONI Johnston and Tiegs.

Amendment.—Subsequent to the publication of the figure of the genitalia
of this species, a number of dissections was made for the purpose of studying the
complex structure of the second segment of the penis and in so doing the filaments
previously stated as absent were found to be present. When drawing Text-fig. 9
(Johnston and Hardy, 1923), they were mistaken for projections of the lobe and
in consequence the lobe illustrated in Text-fig. 9, H, if corrected, should be rounded
and the filaments made to coincide with what is there shown to be the projection
of the lobe. The presence of this filament should also be corrected in the key and
in the descripticn. It is possible that the filaments will also be found (perhaps in
a vestigial or rudimentary state) in other species, so for the character given in
the key “filamerts absent” should read ‘filaments apparentiy absent”’.

Biology.—The seasonal preponderance of certain Sarcophagas in Brisbane
is a well recognized phenomenon. A few species are readily bred in carrion, the
most persistent of which is S. tryoni. It was noticed, however, that for three
months of the year, although adults were captured, they failed to appear in the
carrion. The reason for this non-appearance amongst bred blowflies during June
to August was discovered during experiments upon the pupal period conducted
under uniform temperature control.

Of all the blowflies tested, and at every temperature experimented with, it was
found that the pupal period was consistent in its duration; a formula was worked
out from a chart, and the maximum temperature at which the flies would hibernate
was calculated, the result being contrasted with the winter behaviour of the flies
bred under more natural conditions.

Under normal conditions maggots of S. tryoni deposited on the 9th May,
required 31 to 149 (average normal 133) days to complete their metamorphosis,
or roughly from one to five months. Of this period seven days were spent in the
carrion, five to eight subsequent days were passed before pupation took place, and
from 19 to 136 (normal average 120) days were spent in the puparium. The
number of adults reared in this experiment was 26. The normal average was
based on figures that excluded a small percentage of abnormally quick
developments.

A still more striking effect was produced by maggots deposited a month
earlier, on the 7th April, when again seven days were spent in the carrion, 6 to 24
days elapsing before pupation took place and 17 to 163 subsequent days (well
over five months) passed before the adult emerged, thus taking from 31 to 177
(normal average 165) days to complete metamorphosis, or roughly one month
and nearly six months. The total number of adults bred in this batch was 111.

Further observations elicited the fact that the adults did not survive a cold
night, but succumbed at a temperature of about 55° F., or perhaps higher.

When incubation tests were charted for the various species of Sarcophaga
experimented with, it was found that the probable maximum _ hibernation

BY G.°H. HARDY. 451
temperature varied with the species, that of S. tryoni being the highest, reaching
60° F. During much of the Brisbane winter the day temperatures scarcely rise
above this, except in the sun, and even in the case where a puparium may be
subjected to 65° F. continuously, 22 days must elapse before the adult emerges.
A few hours daily at such a temperature would cause the pupa to develop
exceedingly slowly, extending the three weeks to as many months. On the same
formula at 70° F. the puparium would be found to take 16 days to complete develop-
ment, but above that the increase in temperature causes a much more rapid
development, until at 95° only about 114 days elapse.

There is a record during January of a nine-day pupal period, this being
the minimum recorded, but 11 days seems to be the normal minimum whilst the
absolute minimum according to formula is 10 days. The record of a nine-day
period is possibly an error that should be applied to NS. froggatti which was
frequently bred with the present species.

Sarcophaga misera and S. peregrina are the two dominant winter species
in Brisbane, and are readily bred during that season. Not only have they a
hibernating temperature lower than that of S. tryoni, but also the ultimate
minimum development period is about six and seven days, nor are the adults
so liable to succumb during cold nights.

For 8S. tryoni, the formula is: (c—10) (y—60) = 60. In this x represents
the period in days and y the temperature (Fahrenheit). It will be found
approximately correct for all temperatures from 65° to 88° F., which is the range
over which the temperature experiments were conducted.

The table given covered four years’ research, but irrespective of the years,
the months are given in sequence, and the flies were bred under more or less
natural conditions. This is the only species of the genus with which the year’s
cycle was practically completed. Even in this case one month was missed, namely
October, but an additional and early deposition in November has rectified the
omission to some extent. From this table it will readily be recognized that on the

average some eight or nine generations of S. tryoni occur during a year; the
maximum could scarcely exceed ten and the minimum is seven.
LIFE CYCLE OF SARCOPHAGA TRYONT.
a teue 2 :
ae Jan. Feb. | March| April May || Sept. Nov. Nov Dec.
Date of Deposition. 18 5 6 7 Be alt) 2 15 18
Time spent in carrion ..| 6 6 6 7 7 7 6 6 5
| |
Time _occupied before.
forming puparium after |
leaving carrion =| 1-11 1-6 6-9 6-24 5-8 5-7 2-10 3-6 3-7
Time spent in puparium| 9-14 | 11-15 | 13-16 | 17-163 | 19-136 || 16-18 | 12-1411 13 11-12
| |
Maximum and Minimum |
number of days occu- | |
pied in completing | | |
metamorphosis 56 |) ISPS 20-26 28-29 31-177 31-149 || 29-32 20-29 20-29 20-24
Normal average number) | | | |
of days occupied in | |
completing metamor- | |
phosis Sf Re SA 21 23 28 165 133 | 30 25 25 22
| | |
Number of specimens |
bred aio 50 510 ane 53 85 12 111 26 25 60 7 Wil
| i |

452 AUSTRALIAN AND EXOTIC -SARCOPHAGID FLIES,

S. misera-group.

Synonymy, twberosa-group, duz-group.

The seventeen forms referred to here, agree very consistently in the shape of
the second segment of the penis. The lobe is well indicated with regard to its
position and may even contain a projection of considerable magnitude, whilst the
lateral process is long, usually rigid and pointing more or less horizontally in
direction. The apical process is almost invariably restricted to a minute point
and the filaments project to at least just beyond the apex of the segment, always
being conspicuous.

S. eta conforms to the above characters and therefore is placed here. The
appendages, lobes and filaments are all exceptionally long but in other respects the
species conforms to the misera-group. Bottcher gave the name tuberosa-group to
this conception without defining it and the name was changed later by Parker to
the duz-group but as misera was the first name published of those species considered
here, the name apparently being correctly applied by Johnston and Tiegs, it became
necessary to change the group name once more.

Senior-White sinks NS. luzonensis and S. ceylonensis as variations of S. duz, but
I think this conception is erroneous. Senior-White and Parker differ in their
methods of examining genitalia and would naturally reach different conclusions.
The following list of names are applicable to species of this group:

*misera Walker, 1849. Australia.

aurifrons Doleschall, 1858; preoccupied by Macquart, 1846. See doleschali Johnston

and Tiegs.

*duxz Thomson, 1868, Pacific Islands.

tuberosa Pandel, 1897, Europe, Asia.

harpax Pandel, 1897, North America, Europe, Asia, India, Philippine and Pacific

Islands.

exuberans Pandel, 1897, Europe, ? Africa and North America.

sarracenoides Aldrich, 1916, N. America.

knabi Parker, 1917, Philippine Islands.

luzonensis Parker, 1919, Philippine Islands.

*eta Johnston and Tiegs, 1921, Australia.

doleschali Johnston and Tiegs, 1921 (new name for aurifrons Doleschall, nec.

Macquart).

*shermani Parker, 1923, Canada.

*ceylonensis Parker, 1923, India, Ceylon and Malay Peninsula.

craggi Parker, 1923, India, Africa.

marshalli Parker, 1923, Europe.

*kohla Johnston and Hardy, 1923, Australia.

flavipalpis Senior-White, 1924. India.

scopariiformis Senior-White, 1927, Ceylon.

SARCOPHAGA MISERA Walker. (Text-figures 2, 4 and 5.)

Sarcophaga misera, Johnston and Hardy, Proc. Linn. Soc. N.S.W., 1923, p. 113,
fig. 14.—S. knabi, Senior-White, Rec. Ind. Mus., 1924, xxvi, p. 284 (N. S. Wales
specimens only).

Synonymy.—Senior White refers to some New South Wales specimens sent
to Patton by Froggatt under the name aurifrons. Froggatt used that name
covering at least five species of which S. misera is one and moreover S. knabi to
which Senior-White referred the specimens has not been found in Australia. It is
very evident that the present case is one of misidentification, the identity of two
species being confused owing to the general similarity of the male genitalia.

Structure.—Various drawings illustrate the apex of the last ventral segment
of the male abdomen, showing conspicuously differences in the various species of

BY G. H. HARDY. 453

the misera-group. The drawing of the same part of the typical form is given here
(Text-fig. 4, representing half only), and it will be noted that this does not agree
with that of any other species of the same group published by various authors.

SARCOPHAGA CEYLONENSIS Parker (Text-fig. 6.)

A paratype specimen of this species, compared with S. misera shows a marked
difference in the dentations along the anterior edge of the anterior appendage
(compare Text-figs. 5 and 6). Unfortunately my male specimen of S. dux from
Hawaii has the penis missing, but judging from the results of comparing the other
parts and characters with those of S. misera, it also should differ in detail from
S. ceylonensis. The dentations referred to on N. misera are consistent for several
hundred specimens examined, not one being found to vary to the extent of missing
one of the dentations. Senior-White in discussing the synonymy leaves no doubt
that he suggested ceylonensis was a synonym of S. dux without having made an
attempt to separate the forms on differences in the genitalia, so it would seem this
matter requires further consideration.

SARCOPHAGA KOHLA Johnston and Hardy. (Text-fig. 3.)

Johnston and Hardy, Proc. Linn. Soc. N.S.W., 1923, p. 113, Text-fig. 14.—
S. luzonensis Parker, Bull. Brooklyn EHntom. Soc., 1919, xiv, p. 43. (Specimens
from Queensland only).

In referring to a specimen from Townsville (Q.), Parker drew attention to the
fact that it did not agree with specimens from the Philippine Islands so he placed
it with his S. luzonensis as an atypical specimen. Probably the specimen conforms
to the one described later as S. kohla.

The male has long been known, but only a few specimens were to be found in
the various collections. More recently the species was bred by me from the contents
of decayed shells at Great Palm Island, near Townsville, and both sexes secured
in this manner. Females were also captured on the wing and at the same time
S. beta, tryoni, aurifrons, omega, gamma and misera were captured but none of
these was bred.

Description.—Female: The chaetotaxy is similar to that of the holotype male
except the outer verticals are strongly developed, the frontals have eight bristles
on one side, nine on the other, there are three rows of postoculars. Only two
intra-alar bristles are present and the discal laterals on the first abdominal segment
are reduced so that the second row is represented by one bristle. The legs are
free of long hairs.

The puparium differs from that of S. misera by having protuberances (Text-
fig. 3) around the apical cavity wherein the larval spiracles were situated; seen
laterally this orifice is thus shown to be surrounded by conical projections whereas
on SN. misera (Text-fig. 2), the corresponding line is practically straight. This
character is enough to confirm the specific status of S. kohla indicating that it
cannot be a variation of 8. misera as has been expressed in correspondence that I
have received.

Habitat. Queensland: Great Palm Island, May, 1925. Female allotype and a
male bred from decayed Molluses, and three paratype females captured on the
wing. Other puparia were secured, but failed to develop.

S. peregrina-group.
The three forms referred to here, agree in the structure of the second segment
of the penis. A most suggestive point with regard to the relationship is to be found

454 AUSTRALIAN AND EXOTIC SARCOPHAGID FLIES,

in the nature of the anterior appendage, a large surface of which is formed into a
mass of minute spines. The widely expanding bifid lobe, the shape of the complex
lateral process and the small apical processes are similar in each case, showing a
very close affinity between the species. There are four names standing under this
group:

*peregrinad Robineau Desvoidy, 1830. Australia.

*fuscicauda Bottcher, 1912, Formosa.

*hudsoni Parker, 1923, Ceylon; placed as a synonym of fuscicauda by Senior-White.
*karnyi, n. sp. Java.

i

Text-fig. 7. Male genitalia of Sarcophaga karnyi n. sp. The lettering
conforms to that given by Johnston and Hardy, 1923, namely: A., forceps or
apical part of them seen posteriorly; B., the same lateral view; C., claspers;

D., another aspect of one or both claspers; H., second segment of penis seen

laterally; F., the same ventral view; G., the same or its apex seen posteriorly ;

H., the same, seen anteriorly; J., apex of the lobes seen from other aspects.

a.a., anterior appendage; a.c., anterior clasper; a.p., apical process; (fil.,

filaments; i.p., interior process; k., Knob; l., lobe; lp., lateral process; p.c.,

posterior clasper.

SARCOPHAGA KARNYI, n. sp. (Text-figure 7.)

Specific identity.—Although I unhesitatingly placed this species as a new
form when first received, I now have reason to believe it may have been confused
with S. fuscicauda Bottcher, originally described from Formosa and recorded from
Java and subsequently other places. Mr. Senior-White has supplied S. fuscicauda
from India and this has a very distinctive character differing from that of the
Javanese form. The main sheath of the second segment of the penis is plain
on the rear in S. peregrina and in the present species, whilst that of the Indian
form is provided with a hood projecting rearwards, a character and position for
an appendage that I have not seen equalled in any other Sarcophagid. As it seems
advisable to record these differences under specific names, the Javanese species is
given a new name and it is taken for granted that the Indian form is identical
with that from Formosa, which I have not seen. S. hudsoni Parker is the same as
the Indian fuscicauda as Senior-White pointed out.

Description.—Male: Head with the outer vertical bristles as short as the
postoculars; ten to fourteen frontals; about six facials; about eleven orals; and two

BY G. H. HARDY. 455

rows of postocular bristles. On the thorax two or three intra-alars, the presutural
and prescutellar bristles are present. The abdomen has on the first segment one
row of three discal-laterals and two submarginal bristles; on the second two lateral
submarginals and on the third one median pair and three lateral submarginal
bristles.

The genitalia are remarkably like those of S. peregrina, the forceps being
similar in outline as well as the penis. The anterior claspers are much longer
than the posterior ones and provided with a broad flange on the inner side at the
apex and on the outer side at the base. Anterior appendage with a minutely
spined area that is represented in the figure by a serrated edge; lobe large and
broad with two conspicuous projections; lateral process broad at the base and
produced into a slender projection continuing from its lower edge; apical process
unpaired and short; between the lobes there is a process that evidently represents
the interior process found on S. peregrina, but where it arises from has not been
determined; not only does this process appear to arise from a different position,
but also it differs in shape.

The chaetotaxy of the legs conforms to the general type; scanty long hairs
occur on the posterior femora only.

The female differs from the male in having the second postocular row of
bristles weak or absent; the discal lateral bristles of the first abdominal segment
are more numerous and are arranged in two rows.

Habitat—Java: Buitenzorg; holotype, allotype and a female paratype were
bred from a beetle (Xylotrupes) in June, 1924, whilst the paratypes male
(16.5.1923) and female (11.5.1923) and another female undated were bred from
Gryllotalpa. All these were received from Dr. H. H. Karny after whom the species
is named.

: Other Groups.

Sarcophaga bancrofti and S. fergusoni, two Australian forms, are undoubtedly
allied, as also are S. gamma and S. ochidea judging from Bottcher’s drawing of the
latter. From a comparison of the illustrations I, would have added to the latter
pair S. albiceps Meigen, as being possibly allied but on receiving a consignment
from Mr. Senior-White, from India, I was astonished to find that this form con-
formed to S. omega with regard to type, showing a third relationship and a quite
unexpected one. The alliance of S. synia with S. carnaria has already been
suggested and to these may be added, perhaps, NS. crinata Parker, which in Senior-
White’s drawing appears to have the same shape of penis although this would not
be suspected from Parker’s illustration. I think enough has been given to indicate
that a systematic search for alliances amongst the Sarcophagas for the purpose of
forming groups based on suitable characters and arising from a detailed study of
the genital parts must lead to a great advance in the taxonomic treatment of the
at present rather unwieldly genus Sarcophaga. The following notes deal with the
genitalia of exotic Sarcophagas that have been described in Australian literature,
and the types of which have at some time or other become available to me. These
new notes will make clear some fundamental characters that: the original
descriptions and drawings leave somewhat obscure.

SARCOPHAGA MILLERI Johnston and Tiegs. (Text-fig. 8.)
Rec. Aust. Mus., xiii, p. 185, 1922.

This is the only species yet described from New Zealand. When examining

the type in the Australian Museum, I concluded the genitalia were mutilated, but
E

456 AUSTRALIAN AND EXOTIC SARCOPHAGID FLIES,

from further material received from Dr. R. J. Tillyard, it is evident that what was
taken to be a mutilation was the normal shape of the genitalia. A drawing from
the type is given here where it will be noticed that a large appendage having five
or more digits, claw-like in shape though varying in size, is the leading feature of
the structure. This occurs only on one side, in a corresponding position on the
other side is a short stump giving the appearance of a mutilation. Another
character that is not apparent on the original drawing is the outward directed
apex of the anterior clasper. Much more material is required before the parts of
the penis can be homologized with the corresponding structures in Australian
species. The asymmetrical nature is unique unless, perhaps, this is also to be found
in the next to be described, S. multicolor.

8 E.
A. B

F
oe

Text-fig. 8. Male genitalia of Sarcophaga milleri J. and T. from the type.
Lettering as Text-fig. 7.

SARCOPHAGA MULTICOLOR Johnston and Tiegs. (Text-fig. 9.)

Rec. Aust. Mus., xiii, p. 187, 1922.

Although taken from the same specimen, the free-hand drawing made by Tiegs
and that given here done by aid of the camera lucida, show fundamental differences
in the structure of the forceps and clasper and in the general outline of the penis.
It is scarcely likely that these differences are brought about by slight differences
in the direction of view and so are best accounted for by differences in the
individual conception of the genitalia. The penis is certainly a most complex
piece of mechanism and it is extremely difficult to determine its components. A
few of the parts remind one of the same parts on S. milleri, and were-it not for the
apparently mutilated process being on the opposite side of the penis, I would have

suggested a possible alliance therewith. The two insects are distinct in every
other respect.

BY G. H. HARDY. 457

Text-fig. 9. Male genitalia of Sarcophaga multicolor J. and T. from the type.
Lettering as Text-fig. 7.

Text-fig. 10. Male genitalia of Sarcophaga haemorrhoidalis Fallen, from a
specimen described by Johnston and Tiegs. Lettering as Text-fig. 7.

458 AUSTRALIAN AND EXOTIC SARCOPHAGID FLIES,

Only one of the claspers was detected on the type, and from its position would
appear to be the posterior one. The penis is extremely complicated and a process
present on one side appears to have been broken off from the other; its supposed
shape is indicated by dotted lines in the figures. The species was described from
Rabaul, New Britain, the type being unique.

SARCOPHAGA HAEMORRHOIDALIS Fallen. (Text-fig. 10.)

The Hawaiian specimen described by Johnston and Tiegs has been further
studied and more detailed drawings of the genitalia are incorporated here.

SARCOPHAGA PALLINERVIS Thomas. (Text-fig. 11.)

The genitalia on that Hawaiian specimen illustrated by Johnston and Tiegs has
been drawn in further detail and as it is not easy to homologize the parts given
in the two figures, these are indicated by the letters a, b, c and d. Compare
illustration with those of Aldrich “‘Sarcophaga and Allies’, No. 49, S. peniculata
Parker, and Nos. 120, 121, 8. communis Parker; presumably these, S. floridensis
and S. pectinata, are related.

i

Text-fig. 11. Male genitalia of Sarcophaga pallinervis Thomas, from a specimen
described by Johnston and Tiegs, Lettering as Text-fig. 7.

BY G. H. HARDY. 459
List of Works Referred to.

JOHNSON AND Harpy, 1923.—Proc. LINN. Soc. N.S.W., xlviii, pp. 94-129. This incorporated

a list of eighteen works to which must be added for the purposes of the present paper
PARKER, 1914.—Proc. Boston Soc. Nat. Hist., xxxv, pp. 1-77.

, 1917.—Proc. U.S. National Museum, liv, pp. 89-97.
, 1919.— Bull. Brooklyn Entom. Soc., xiv, pp. 41-46.
, 1923.—-Ann. Mag. Nat. Hist. (9), xi, pp. 123-129.

SENIOR-WHITDP, 1924.—Rec. Indian Mus., xxvi, pp. 193-285.

, 1927.—Spolia Zeylanica, xiv, pp. 77-83.

NOTES ON AUSTRALIAN MARINE ALGAE.
iv. Tue AUSTRALIAN SPECIES OF THE GENUS SPONGOCLONIUM.
Bye AG Ee Ss GwGAS I MEASIBAS Cs

(Plates xxvii-xxxv.)
[Read 28th September, 1927.]

In his Synoptic Catalogue of Australian and Tasmanian Algae, published in
the concluding volume of Phycologia Australica, 1863, Harvey ranked fifty species,
divided into seven sections, under Lyngbye’s genus Callithamnion. In De Toni’s
Sylloge Algarum only nine of these are retained in Callithamnion. The rest have
been drafted into the genera Chantransia, Spermothamnion, Monospora, Spongo-
clonium, Warrenia, Antithamnion, Lasiothalia and Thamnocarpus.

Harvey defined his Section 1, Dasythamnion, as including species with rather
large fronds, stupose-spongiose, the stem and branches more or less covered with
a veil of decurrent threads, intricate, free and anastomosing. The plants of
this section have been divided between two genera, Spongoclonium and Lasiothalia,
the former ecorticate and the latter corticated. Both have the characteristic
cystocarps and sporangia of Callithamnion.

The history of the two genera has been somewhat remarkable. Both were
proposed in 1854 almost simultaneously, so that it is apparently impossible to
decide which may claim priority. Spongoclonium was proposed by Sonder,
Linnaea xxvi, p. 515 on a specimen which he called S. conspicuum, and Lasiothalia
by Harvey, Trans. Roy. Irish Acad., xxii, p. 558, “founded on a very imperfect and
battered scrap” (Harvey), which he called L. hirsuta.

In the Flora of Tasmania, 1860, Harvey gives a description of Callithamnion
conspicuum Harv. and appends the comment: ‘Since this was written, I have
ascertained that my C. tingens is the same as Sonder’s Spongoclonium conspicuum,
which name will be adopted if the plant be removed from Callithamnion. Another
synonym is my Lasiothalia hirsuta, founded on a very imperfect and battered
scrap of what I afterwards called Callithamnion tingens when more perfect
specimens were collected”. Presumably he called the species C. conspicuum Harv.
because he as yet retained it in Callithamnion.

Of his C. conspicuum, which we see he identified with Sonder’s plant, Harvey
writes in the Flora of Tasmania: “The habit of this species is peculiar, and
perhaps, with C. plumigerum and C. superbiens, it might form a separate genus,
characterized by the peculiarly inarticulate and hirsute stem. But there are
many intermediate links with species of more ordinary characters”.

Sonder seems to have acquiesced in the inclusion of his species in
Callithamnion, as in his enumeration of Australian Algae he drops Spongoclonium
and lists his plant as (. conspicuum.

It might seem then that the matter was definitely settled, that but one species
existed, and that the two authors had withdrawn their genera, leaving the plant

BY A. H: S. LUCAS. 461

in question in the mother genus Callithamnion. But J. G. Agardh (Analecta
Algologica, 1892) voiced his doubt of the identity of the species. His grounds
appear somewhat slender. He says that Harvey, writing of Lasiothalia, says that
the plant but slightly adheres to paper; Agardh’s specimens (probably distributed
by Harvey as OC. tingens) adhered so closely that he could scarcely
separate the least fragment, and he suggests that Harvey’s name hirsuta of itself
seemed to point to another plant. De Toni, in Sylloge Algarum, iv, 1903, on the
other hand, regards S. conspicuum Sond., C. conspicuum Harv., C. tingens Harv.
and Lasiothalia hirsuta Harv. as being synonymous, and, acting on Harvey’s
suggestion given above, adopts the Harveyan genus Lasiothalia incorporating in
it L. hirsuta and the three Harveyan species ©. plumigerum, C. formosum and
C. superbiens.

In 1892 (Analecta Algologica) Agardh resuscitated Sonder’s genus Spongo-
clonium, incorporating in it the MHarveyan Callithamnion brounianum,
C. wollastonianum, OC. dasyurum, C. formosum, and adding a new _ species,
S. wilsonianum, but curiously left Sonder’s type species amongst the Callithamnia
as C. conspicuum. Ii then, as Agardh thought, S. conspicuum and L. hirsuta are
not identical, the former has found no resting place in the Sylloge Algarum.

If the type species of a genus, the only species recognized by the author of the
genus, is removed from it, it would seem that the genus lapses. Agardh does
not seem to be justified in remodelling the genus without including in it
S. conspicuum. _He considered that C. tingens was a different plant, but still did
not retain S. conspicuwm. From material which I have gathered I believe that
Agardh was right, that there are two species, that C. tingens is distinct from
Sonder’s plant, and I propose to retain S. conspicuum with his description in the
genus Spongoclonium Sonder, the retention of which is thus justified.

In 1894 (Analecta Algologica contd.) Agardh took into Spongoclonium
Harvey’s C. plumigerum and added two new species of his own, S. fasciculatum
and S. scoparium. R. M. Laing (Trans. N.Z. Inst., xxxvii, 1905) contributed two
fresh species from New Zealand, S. pastorale and S. brachygonum.

In Sylloge Algarum, iv, 1903, De Toni enlarged the genus Spongoclonium by
adding to it the Harveyan C. latissimum,* C. angustatum, C. hirtum* (from New
Zealand), C. violaceum,* OC. dasytrichum* (from Rio de Janeiro), C. scopula,
C. paradoxum,* C. crispulum* and C. debile,** together with Suhr’s C. stuposum’*
(from the Cape of Good Hope). The species bearing the asterisk he marks as
doubtful Spongoclonia. At the same time he removed 8S. plumigerum and
S. formosum from the genus transferring them to Lasiothalia.

The critical distinction between Spongoclonium and Lasiothalia seems to be
that the former is monosiphonious, articulated, without any true cortex, and
thus nearer to Callithamnion, while the latter possesses a true cortex, and is
nearer to Crouania.

The object of the present paper is to illustrate as far as material will permit
the Australian and Tasmanian species included by De Toni in the genus
Spongoclonium.

It must be admitted that of several of the species the material for investigation
is scanty. Some are not represented in any of the Australian Herbaria and others
but slightly. Many occur but rarely, and have eluded diligent search in the
localities whence they have been recorded. The paper aims to set forth our
present knowledge so that the gaps can be clearly recognized. It is based on
the material existing in the Melbourne and Sydney Herbaria and that which I
have accumulated from the coasts of Victoria and Tasmania.

462 NOTES ON AUSTRALIAN MARINE ALGAE, iV,

I wish to acknowledge with gratitude the courtesy and kind help which
Messrs. Audas and Morris, of the Melbourne Herbarium have shown me, and to
thank the Curators of the Melbourne and Sydney. Herbaria for permission to
publish photographs of specimens in their Herbaria.

SPONGOCLONIUM Sonder.

“Frons teres, rosea, spongiosa, pinnatim decomposita, contexta filis articulatis
callithamnoideis ramosis anastomosantibus, a tubo centrali articulato egredientibus,
exterioribus vel periphericis laxis secundatim ramuilosis ramulis incurvis liberis.
Sphaerosporae sphaericae triangule quadridivisae, in ramulis periphericis latere
interiore evolutae, solitariae vel seriatae, pedicellatae, pedicello simplici vel
subdiviso.

“Genus habitu fere Ptilocladiae, sed fronde tercti molliori non regulariter
distiche pinnata, praeterea filis periphericis liberis subincurvis secundato-ramulosis
non ut in Ptilocladia densis fastigiatis stratumque periphericum efficientibus
diversum. Ab Haloplegmate, cui filis exterioribus liberis et substantia molli
spongiosa affine, facile distinguitur frondi tereti pinnata et axi centrali articulato”.
Sonder, Linnaea, xxvi, p. 515, 1854.

Agardh (Analecta Algologica, p. 39, 1892) gives this amended description of
Spongoclonium.

“Frons articulata monosiphonia, initio et superne nuda alterne pinnatim
decomposita, mox filis a ramorum basi extra cuticulam secus frondem descenden-
tibus subfuniculariter contortis et anastomosantibus inferne stuposa, stupa nunc
fere nuda, nunc hirta, nunc dense ramellis cooperta et spongiosa. Favellae ad
apices ramulorum subterminales, involucro ramellis plurimis incurvatis constante
circumcirca obtectae, plures circa rachidem tabescentem dispositae, singulis
saepe bracteatis, in eundem nidum conjunctae; nucleis minoribus rotundatis
gemmidia plurima rotundato-angulata intra periderma hyalinum foventibus.
Sphaerosporae interiore latere ramulorum paucae (aut saepe plures seriatae),
a singulis articulis transformatae, triangule divisae. :

“Quale sub-genus Dasythamnii inter Callithamnia limitibus circumscripsit
Harvey, genus autem proprium nomine Spongoclonii a Sondero institutum fuit’’.

Here we have the idea of an alternately pinnately decompound Callithamnion,
which covers the axis of its stem and branches to a variable height with a
growth of descending radicules which interweave into a close tegumentum; the
cystocarps protected by an involucre of several small incurved branches; the
sporangia, few or seriate, divided into tetrads.

Agardh has added the character of the cystocarps, and does not postulate
that the sporangia be pedicellate.

We now give our working definition in the light of further investigation and
so as to include NS. conspicuum.

SPONGOCLONIUM Sonder.

Sonder, Linnaea, xxvi, p. 515, 1854; J. G. Agardh, Anal. Algol., p. 39, 1892;
Engl. and Prantl, Natiurl. Planzenfam., p. 491, 1897; De Toni, Sylloge Algarum, iv,
Dp) 135771903:

Frond erect, rounded, with alternately pinnate primary branches of unequal
length, spreading in all directions; these bear similarly ramulose secondary
branches, the whole frond being thus pinnately decompound. The lower cells of
the branches emit much branched radicules, which becoming decurrent clothe
the main stem and branches with a spongy network, at all events in the inferior

9

BY A. H. S. LUCAS. 463

part of the plant. A central monosiphonious articulated axis is continuous
throughout the stem and branches, ecorticate but invested below by the descending
radicules and their branches.

Cystocarps numerous, scattered over the outer filaments of the frond, situated
outside the close reticulum among the looser ramelli but with no proper involucre.
The nucleus consists of a single gonimoblast divided in successive evolution into
numerous globose-rounded few-celled gonimoblasts.

Sporangia shortly pedicellate or sessile, few or seriate, growing on the inner
side of penultimate or ultimate filaments, divided into tetrads.

Antheridia thyrsoid.

Sections.
The species fall naturally into two sections with quite different aspects.
Section 1. Conspicuae.

The stem and greater part of the branches stupose-spongiose, formed of
intricate funicular descending radicules, only the terminal plumules free and
articulate.

. conspicuum.

. fasciculatum.

. brounianum.

. wollastonianwm.
. wilsonianum.

. dasyurwm.

. paradoxum.

RARRRRAR

Section 2. Violaceae.

Only the lower part of stem and branches funicularly intricate, the greater
portion of the frond free and articulate.

S. latissimum.
S. angustatum.
S. violaceum.

S. scopula.

S. scoparium.

We propose to exclude from the genus and to restore to Callithamnion the
species—
S. (€?) crispulum (Harv.) De Toni.
S. (2?) debile (Harv.) De Toni.

Section 1. CONSPICUAE.
S. CONSPICUUM Sonder.
Linnaea, xxvi, p. 515, 1854.

Specimen 4 inches high, lacking the attachment. Primary frond terete,
half a line wide, bipinnately branched. Branches for the most part distichous,
mingled with some arising irregularly from the frond, subhorizontally patent,
narrower by half than the caulis, the lower about an inch long, the upper longer,
about 2 inches in length, all alternate with pinnae, 2-4 lines long. Sphaerospores
on the inner side of subarcuate filaments, now solitary, now longitudinally seriate;
on a short 1-2 jointed or a longer 4-6 jointed, simple or sub-branched, pedicel,
spherical, the nucleus triangularly divided. Frond spongy, rose-sanguineous. <A
central axis monosiphonious, articulate, persisting through the whole frond, at
the genicula beset with callithamnoid branching and anastomising threads, free
towards the surface. Joints of the central axis 3-4 times as long as the diameter,
of the ramelli twice as long.

Habitat—Cape Liptrap, June, 1853.

G

464 NOTES ON AUSTRALIAN MARINE ALGAE, iv,

S. FASCICULATUM J. Ag. (Plate xxvii.)
Anal. Algol. contd., p. 118, 1894.

Frond elate, to 26 cm., stout, caulescent, pinnately decompound; primary
branches alternate at distances of about 5 mm., pyramidal, rather patently
ascending; branches of the second order, pinnae, 20 mm. long, emitted alternately
at rather shorter distances, similarly patently ascending; ramuli of the third order,
soft, fasciculate, with capillary ramelli, the ultimate branches of which are
somewhat distant, ultra-capillary and but slightly diverging. The stem, primary
and secondary branches are densely clothed by the intricate web of the ramelli
which they emit. The secondary branches incurve towards the axis on which
they grow.

The joints of the upper parts of the axis are 2-4 times as long as the
diameter, those of the felted ramelli are often much shorter, even falling to half
the diameter.

The colour of the fascicles is bright rose, of the felt covered stem and branches
rusty brown.

Cystocarps “several among the: fascicles and confluent into a conspicuous
node” (J. Agardh).

Sporangia not seen.

Antheridia numerous, oblong, sessile, borne on the upper branches of the
ramelli.

Habitat.—Port Phillip Heads and Western Port (Wilson). Apparently very
rare. The antheridia-bearing specimen described and figured is in the Melbourne
Herbarium. It is the only example we possess in Australian Herbaria.

S. BROUNIANUM (Harv.) J. Ag. (Plates xxviii and xxix.)

Callithamnion brounianum Harv., Trans. Roy. Irish Acad., xxii, p. 561, 1854.

Frond elate—in the Western Australian forms, first described by Harvey,
to 10 cm., in those afterwards obtained by him from Western Australia and Port
Fairy to 30 cm.—caulescent, pinnately decompound; primary branches alternate,
spreading, to 10 cm. long, with lateral divisions often as thick as the main axis;
secondary branches arising from these or from an undivided primary, to 2 cm.
long, bearing alternately at very short intervals the fascicles of the last order.
The stem and primary and the basal parts of the secondary branches densely
stupose, invested by a close felt of squarrose-pilose alternately pinnate ramelli
derived from descending radicular branches. A continuous ecorticate articulated
axis persists through stem and branches. The joints in the lower portions about
as long as the diameter, in the ramelli very variable, short to twice the diameter.
The divisions of the ramelli copious, pinnate, capillary, the upper incurved.

Colour of the plumules deep rose red, of older axes paler and browner.

Cystocarps “nidulating among the ramelli below the corymbs” (De Toni).

Sporangia numerous, seriate on the inner side of the lower parts of the ramelli,
very shortly pedicellate or almost sessile, with clear spherical envelope, dividing
into tetrads.

Habitat.—Fremantle (Clifton), King George’s Sound, Port Fairy, Queenscliff
(Harvey), Port Phillip Heads (Wilson).

Cystocarpiferous plants seem to be rare.

R. M. Laing, Trans. N.Z. Inst., xxxvii, pp. 392, 393, 1905, describes a form
Pleonosporium brounianum Harvey Gibson from New Zealand, which he identifies
with the Australian Callithamnion brounianum Harvey. In the New Zealand
plants the sporangia are pear-shaped and develop 8, 16 or 32 radially arranged

BY A. H. S. LUCAS. 465

spores, and hence Laing transfers the species to the genus Pleonosporium. In
our plants the sporangia are spherical, and when mature divide each into a tetrad
of spores. This I have fully verified in specimens distributed by Harvey in his
Algae Australicae Exsiccatae. Laing also places the New Zealand species C. hirtum
H. and H. in Pleonosporium from the sporangia, and thinks that the tetrad
sporangia figured by Harvey in Flora Antarctica, t. 78, are explained as being
derived from some other plant. It would seem then that Spongoclonium
brounianum is not known to occur in New Zealand, and that S. hirtum is really
a Pleonosporium.

S. WOLLASTONIANUM (Harv.) J. Ag.

Callithamnion wollastonianum Harv., Trans. Roy. Irish Acad., xxii, D. 561, 1854.

No example of this plant exists in any of our Australian Herbaria. The
following is derived from Harvey’s original description, supplemented from the
description in the Sylloge Algarum.

Frond elate, caulescent (according to Harvey, ultrasetaceous), 10 to 14 cm.
high, thinly corticate at the base, the stem for a long distance upwards stupose-
hairy with squarrose pile derived from radicules descending from the basal joint
of superior branches, sub-distichously freely branched, the branches alternately
Pinnately decompound, those of the last order distichous, pellucidly articulate,
alternately plumulate, the plumules very long, linear in outline; the pinnae
slender patent (rather erect, Harvey) short, the inferior simple, the superior more
often forked or pinnulate. The joints 2-4 times as long as the diameter (Harvey),
of the branches 13 times (De Toni, Sylloge).

Cystocarps and Antheridia not noted.

Sporangia solitary along the ramelli, very shortly pedicellate.

Habitat.—Middleton Bay, King George’s Sound, rare (Harvey).

S. WILSONIANUM J. Ag. (Plate xxx.)
Anal. Algol., p. 42, 1892.

Frond elate, 10 to 15 cm. high, caulescent, alternately pinnately decompound;
primary branches spreading in all directions, to 10 em. long; secondary emerging
at short intervals, patent, distichous, plumose, lanceolate in outline, to 15 mm.
long, bearing tertiary alternate, distichous, plumulose ramelli, 1 or 2 mm. long;
the stem and rachides of the primary and even of the secondary branches densely
clothed with an intricate felt of the segments of ramelli of slender descending
radicules; a continuous ecorticate monosiphonious axis persists through stem and
branches, the joints throughout about equalling the diameter in length, the
copious divisions of the ramelli all alternately pinnate, capillary, the upper
incurved.

Colour a dull red, of older rachides brown.

Cystocarps and Antheridia not observed.

Sporangia numerous, nidulating within the free plumules, borne laterally on
the lower joints of the ramelli, subpedicellate, spherical, divided into tetrads.

Habitat—Port Phillip Heads (Wilson).

It will be seen that these four species conform to Sonder’s conception of the
genus, that they are all very closely related and form a very natural group.
Harvey was at first doubtful whether or not to include the smaller and the larger
plants he found in the one species S. brownianum, but finally combined them.
Of the other three very few examples have been found, S. fasciculatum and

466 NOTES ON AUSTRALIAN MARINE ALGAE, iv,

S. wilsonianum only in Wilson’s dredgings, while of S. wollastonianum we have
no specimen in Australian Herbaria.

The habit of S. fasciculatum differs from that of the others, the branches in
these are spreading, in S. fasciculatum distinctly ascending. The terminal ramuli
in the latter are fasciculate, and the segments more finely capillary and longer
and more erect and possess longer joints. S. wilsonianum is more delicate than
S. brounianum, with feather-like plumules in contrast with the more brush-like
pinnules of S. brounianum. Of S. wilsonianum De Toni remarks that it is very
near S. wollastonianum but larger and in every respect more evolved. I take
the latter to mean more decompound.

S. pasyurum (Harv.) J. Ag. (Plate xxxi.)
Callithamnion dasyurum Harv., Synoptic Catalogue, p. 51, 1863.

Frond elate, to 22 cm. high, alternately distichously pinnate-decompound, the
pinnae and pinnules ascending virgate, most of the plant spongiose-stupose, with
decurrent interwoven threads and even the smaller ramelli in the same way hairy,
the pinnules scarcely stupose alternately closely plumulate, the plumules minute
oblong pectinate-pinnate. Their pinnellae elongate incurved, simple or ramuliferous
at the base; all joints very short. Colour intensely red.

Cystocarps, Sporangia, Antheridia not seen.

Habitat.—Port Phillip Heads (Harvey).

There are no specimens in the Melbourne Herbarium, and but one in that of
Sydney, Harvey’s No. 508E. This, the one figured, gives the appearance of a much
denuded plant, the plumules being very weakly developed, and adhering so closely
to the paper that it was impossible to dissect them under the microscope.

S. PpARADOXUM (Harv.) De Toni. (Plate xxxii.)
Callithamnion paradozum Harv., Flora of Tasmania, ii, p. 337, 1860.

Frond terete, decompound pinnate, branches and branchlets spreading in all
directions, the main branches pyramidate in contour, the rachides stout below,
gradually attenuate, all except the ultimate plumules densely spongy, formed of
closely interwoven descending filaments; the surface of the frond consisting of
the crowded pinnae of penultimate order, the lower branches of these greatly
elongated, the upper markedly flexuose with distichous segments ending in blunt
apices. The joints are three or four times longer than the diameter in the lower
part of the frond, not much longer than the diameter in the ultimate free ramelli.

Colour.—A rich dark plum colour.

Cystocarps and Antheridia not seen.

Sporangia numerous, sessile, seriate on the upper or inner side of the ramelli
near the surface of the plant, rather large, 62 uw in diameter,

Habitat—Warrnambool (H. Watts), Sandringham, Port Phillip (Lucas) ;
Tasmania, Brown’s River (Gunn), estuary of the Derwent River and Orford
(Lucas).

It is very abundant in the Lower Derwent. I have gathered many
sporangiferous plants but was never fortunate to discover cystocarps.

De Toni placed this species in the genus with some doubt. It seems to me to
fall under Sonder’s definition admirably. It is the most clearly marked of all
the species.

BY A. H. S. LUCAS. 467

Section 2. VIOLACEAE.
S. LATISSIMUM (H. and H.) De Toni. (Plate xxxiii.)

Callithamnion latissimum H. and H., London Journal of Botany, iv, p. 452,
1845.

Frond elate, caulescent, spreading pinnately decompound, 8 to 12 or even to
25 em. high and to equally broad; primary branches alternate, beset with very
slender, repeatedly divided pinnae; the stem and main branches, primary and
secondary, far upwards stupose-hairy from radicules descending from the basal
joint of the next superior segment; a monosiphonious axis is continuous through-
out the branching forming the design of the plant, entirely ecorticate but
surrounded below by an intricate investiture of the radicules and their branches;
all the free tertiary pinnules plumulose, much and finely divided, the ramifica-
tions throughout alternately pinnate.

Colour of the plumules rosy in young and sterile plants, duller and even a dark
purple in fruiting plants, of the felted rachides brown.

Cystocarps single, terminal on short segments, involucrated by inferior
incurved ramelli.

Sporangia spherical, about 35 uw in diameter, numerous, distinctly pedicellate,
the pedicel unicellular nearly or quite as long as the diameter of the sporangium,
often subseriate secund on the inner side of the ramelli, forming tetrads.

Antheridia not observed.

Habitat.—Tasmania, especially in the Tamar River (Gunn, Harvey, Fereday,
Lucas), Port Phillip (Wilson, less luxuriant, Lucas).

S. aneustatum (H. and H.) De Toni. (Plate xxxiv, fig. 1.)
Callithamnion angustatum H. and H., Flora of Tasmania, ii, p. 334, 1860.
Frond elate, to 12 cm. high, pinnately decompound, below funicularly contoried

and somewhat stupose with radicules descending the rachis from the basal joints
of the larger branches, and above and in greater part emitting free and very
flaccid branches; the branches soft, in all directions pinnate, the pinnae, on an
elongated axis with rather naked protruding apex, alternate, conspicuously
attenuated from the base, the lowest pinnae of the larger branches subrecurved,
the upper short patent; the joints four times as long as the diameter.

Colour a bright rose-red, brighter and lighter than in S. latissimum.

Cystocarps geminate, involucrated by subumbellate ramelli of unequal length.

Sporangia scattered on the ramelli, quite sessile, divided into tetrads.

Antheridia not observed.

Habitat.—Tasmania, Georgetown (Harvey), Tamar _ River, Ulverstone,
D’Entrecasteaux Channel (Lucas), Port Phillip Heads (Wilson); grows on other
algae.

The slender stem and primary branches are usually quite inconspicuous, and
hence difficult to trace amidst the superabundance of ramuli of secondary and
higher orders. In a cystocarpiferous plant from Ulverstone, however, they were
as conspicuous as in the upper regions of S. latissimum. The larger branches are
funicularly contorted and give off radicules, the radicule now passing down the
stem from the basal joint of a branch, now formed from the lower pinnules of the
branches. :

The cystocarps opposite geminate, on a smaller ramus below the persisting
apex, are surrounded by a sort of involucre of subumbellate unequal ramelli,
several of the latter acuminate with shorter joints.

The habit is quite that of Callithamnion.

468 NOTES ON AUSTRALIAN MARINE ALGAE, iv,

S. vioLacEuM (Harv.) De Toni. (Plate xxxiv, fig. 2.)
Callithamnion violaceum Harv., Flora of Tasmania, ii, p. 334, 1860.

“Caespitose, purple, fronds capillary, stupose at the base to a considerable
height with radicular threads interwoven into cords, pellucidly articulate decom-
pound pinnate; the branches in all directions pinnate at the base bipinnate at
the apex, pinnate with simple plumules, pinnules patent filiform elongate; joints
of the rami six times, of the ramuli three times, as long as the diameter;
tetraspores sessile on the pinnules, subsolitary, globose.

“A small species 2-3 inches high. The principal branches are bundled together
into ropes, and then closely interwoven by root-like fibres, which issue from the
nodes, and proceed downwards along the stem, forming an accessory stupose
stratum. The branches are alternately pinnate below, and bipinnate above, all
the pinnules remarkably patent and elongate. Tetraspores are thinly scattered on
the inner faces of the pinnules’’.—(Harvey.)

Colour a dull purple.

Cystocarps geminate with no involucre of ramelli (Lucas).

Antheridia not observed.

Habitat.—Tasmania: Georgetown (Gunn, Harvey), abundant in the Derwent
Estuary (Lucas).

Specimens found by Gunn, much larger, with stronger and shorter ramuli of
rather shorter joints, may, in the opinion of De Toni, constitute a distinct species.
I have had no opportunity of seeing Gunn’s specimens, nor have I gathered forms
of the kind.

'§. scoputa (Harv.) J. Ag.

Callithamnion scopula Harv. -Trans. Roy. Irish Acad., xxii, p. 562, 1854.

Frond small, an inch high, below funicularly contorted and rooting, at length
stupose at the base, above terminated in free fascicles of branches, the greater
branches gradually on all sides porrect pinnate, the pinnules subdistichous rather
simple elongate, curved, the lower subhorizontal recurved, the upper patent, joints
23 times as long as the diameter.

Cystocarps and Antheridia not seen.

Sporangia elliptical, sessile, scattered among the pinnules.

Habitat.—Rottnest Island, Fremantle (Harvey).

We have no specimens in Australian Herbaria, but the plant would appear to
be a smaller Western Australian representative of the Tasmanian S. violacewm.

S. scoparium J. Ag.

Anal. Algol. contd., i, p. 117, 1894.

Frond caespitose-rounded, expanded in every direction, composed of branches
repeatedly fasciculately divided, the branches radiating above a stipes less
conspicuously funicularly compound, branches and branchlets issuing on all sides
erect-patent, rather rigid, of unequal length, joints slightly contracted at the
dissepiments usually three or four times as long as the diameter. No fruits seen.
The bushy fronds stout, measuring 10 cm. in expansion.

Habitat.—Western Port (Wilson), Orford, Tasmania (Mrs. Meredith).

We have no examples in the Australian Herbaria. There seems but little in
this meagre description to separate the species from NS. violacewm from the same
region. Is it the same thing as Gunn’s larger form of S. violaceum?

BY A. H. S. LUCAS. 469

To be excluded from the genus Spongoclonium.
CALLITHAMNION DEBILE Hary. (Plate xxxv, fig. 1.)

Trans. Roy. Irish Acad., xxii, p. 563, 1854; Spongoclonium ?? debile (Harv.)
De Toni, Sylloge Algarum, iv, 1908.

Frond small, caespitose, scarcely an inch high, ultracapillary, decompound
pinnate, the lower branches giving off ramuli on all sides, the upper distichously
pinnulate, the pinnules very patent, inferior articuli 5-8 times, and those of the
ramuli 3-4 times, as long as the diameter.

Colour purpurascent.

Cystocarps and Antheridia not seen.

Sporangia solitary, sessile on the ramuli (Harvey).

Habitat.—Rottnest Island, Fremantle (Harvey), “In shady crevices of rocks
at half-tide level”.

There are specimens in the Melbourne and Sydney Herbaria, distributed by
Harvey. The specimen figured, named by Harvey, was gathered by himself at
Geelong. De Toni states that he had not seen authentic plants. Examples of the
genuine Western Australian plant are needed.

CALLITHAMNION CRISPULUM Harv. (Plate xxxv, fig. 2.)

Trans. Roy. Irish Acad., xxii, p. 562, 1854; Spongoclonium (?) crispulum
(Harv.) De Toni, Sylloge Algarum, iv, 19038.

Frond small, caespitose, 1-3 cm. high (half to three-quarters of an inch,
Harvey), decompound pinnate, with few pyramidate capillary branches bearing
short crowded pinnules, the whole ecorticate and nowhere spongy, but with simple
Callithamnion structure. From the stem, and from the same articulus, in a very
few cases a branch proceeds upwards and a simple elongate free arcuately curved
radicule hangs downwards. These radicules at first of the colour of the plumules,
and with joints of the same length, become hyaline as they descend and the joints
become longer. The plumules are alternately distichously pinnulate, the pinnules
simple, elongate, incurved, the lower patent, the upper corymbose, the joints one
and a half times as long as the diameter.

Colour dull red.

Cystocarps geminate (Harvey).

Sporangia and Antheridia not seen.

Habitat.—Rottnest Island, Fremantle (Harvey). Both Melbourne and Sydney
Herbaria possess specimens distributed by Harvey.

The presence of occasional free radicules does not at all seem to warrant the
inclusion of this well marked plant in Sonder’s Spongoclonium, and it would, I
think, be best left in Callithamnion.

EXPLANATION OF PLATES XXVII-XXXV.
Plate xxvii.
Spongoclonium fasciculatum. Bearing antheridia. "Western Port (Wilson). Melbourne
Herb.
r Plate xxviii.
Spongoclonium brounianum. Bearing tetrasporangia. Port Phillip Heads (Wilson)
Melbourne Herb.
Plate xxix.
Spongoclonium brounianum. Fremantle (G. Clifton). Melbourne Herb.

470 NOTES ON AUSTRALIAN MARINE ALGAE, iv.

Plate xxx.

Spongoclonium wilsonianum. Port Phillip Heads (Wilson). Melbourne Herb.

Plate xxxi.

Spongoclonium dasyurum, Harvey, Alg. Austral. Hxsice. Sydney Herb.

Plate xxxii,

Spongoclonium paradoxum. Orford, Tasmania (Lucas).

Plate xxxiii.

Spongoclonium latissimum. Tasmania. Melbourne Herb.

i)

bo

Plate xXxXxiv.
Spongoclonium angustatum. Georgetown, Tasmania (Harvey).
Spongoclonium violaceum. Derwent River, Tasmania (Lucas).
Plate xxxv.

Callithamnion debile. Geelong (Harvey). Melbourne Herb.
Callithamnion crispulum. Fremantle (Harvey). Melbourne Herb.

STUDIES IN THE GOODENIACEAE.
I. Tue Lire-History oF DAMPIERA STRICTA (R. Br.).

By P. BroucH, M.A., B.Sc., B.Sc. (Agr.),
Lecturer in Botany, the University of Sydney.
(Plates xxxvi-xxxvii and fifty-eight Text-figures. )
[Read 28th September, 1927.]

Introduction.

One of the final attainments of botanical research will be the realization of an
absolute Natural System of Classification of plants.

How far this goal is from attainment is manifest, and even more apparent
is the difficulty of such an achievement. The nature of the problem does not
permit of its being resolved by the brilliancy of the individual. The arrangement
of plants in such a manner as to show their phylogenetic relationships can be
achieved only through the cumulative results of researchers throughout the ages.
The magnitude of the task is colossal, but the lure of it is a splendid incentive to
effort, and each worker may contribute his mite.

No phylum has received so much attention as the Angiosperms—a fact
readily understood—yet, despite the efforts of systematists from the time of
Linnaeus to the present day, the so-called Natural System of Classification of
Angiosperms, is still largely artificial, and a matter of surmise. The Orders and
even many Families are, in the main, merely aggregations of forms with similar
gross characters, that is, are form groups, which may or may not consist of
closely related members. The variety of Systems, which claim precedence at the
present day, bears testimony to the above statement. Agreement has not even
been reached as to what constitute the most primitive forms of flowering plants.

It would seem that further progress may most confidently be expected along
that line of investigation which deals with the life-histories of representatives of
those Families about which little or nothing beyond gross morphology is known.
In this way assumptions or suspicions may be tested, and unsuspected relationships
exposed.

For obvious reasons our knowledge of the life-histories of Angiosperms is
largely confined to cosmopolitan forms, or to those prevailing in the Northern
Hemisphere, and particularly to the Temperate Regions. It is the more desirable
then to gain information regarding representatives of floras typical of the great
land masses south of the Equator, such as South Africa, South America and
Australasia. Most particularly does this apply to Australia, which, on account of
its long isolation from adjacent continents, may well harbour forms of peculiar
significance. In addition the study of endemic forms may shed light on the vexed
question of the original centre of distribution of Flowering Plants.

So far, very little indeed has been done along such lines in Australia, which

with a flora noted for its high percentage of endemism may supply some interesting
information concerning the course of evolution.
H

472 STUDIES IN THE GOODENIACEAE, 1,

Only two Families of plants are confined to Australia, viz. Brunoniaceae and
Tremandraceae, but at the same time many Families find their chief expression
in Australia, and beyond its borders are represented only by a few widespread
species. An outstanding example of this is furnished by the Goodeniaceae, a
Family which has many points of interest, but which is chiefly noteworthy on
account of its indusiate stigma, and peculiar pollinating mechanism.

Such abnormalities, coupled with the puzzling distribution of the genera
and species comprising the Family certainly held the attention of the writer, and
suggested that an investigation of the life-history of a representative of one or
more genera might not be an unprofitable undertaking.

THE GOODENIACHAE.

The Goodeniaceae comprises 12 genera which have been divided into two
sections (Bentham and Hooker, 1869).

Section A. Leschenaultia, Anthotium, Velleia, Goodenia, Calogyne, NSelliera,
Catosperma—consisting of forms with two or more ovules in each loculus of the
ovary, or on each side of the imperfect or rudimentary dissepiment.

Section B. Scaevola, Diaspasis, Verreuxia, Dampiera, Brunonia—having one
or two ovules in each ovary.

Some authorities prefer to exclude Brunonia from the Goodeniaceae and
recognize a Family—the Brunoniaceae—which is monotypic.

Engler and Prantl (1887) subdivided the Goodeniaceae into I. Goodenioideae
(including all the above genera except Brunonia) and II. Brunonioideae with a
single genus Brunonia.

An investigation of the life-history of Brunonia is obviously to be desired.

Of the twelve genera quoted nine are confined entirely to Australia (Bentham
and Hooker, 1869), viz.:

Goodenia Dy erie Ete WLC OO S19 105
A IDAMpPlEray a ei sw ce se eae Pe Oho,

Leschenaultia Me ia Pama ee eliG
Vielleiaiy cic.) wiih ie a. aerek Cae SARS
IVETE Use ain eemrcan foveal 35 2
ANTNBOUOUEI Mbi5 05 Ube Vere Bs 2
Catosperma A AeA ex dooge Me nasa
Diaspasis 6a Lee ~ Te
Brunonia seal eatys adage hoe: eu oS 1

while of the remaining three genera, Scaevola with 65 species is distributed
throughout Australia, Polynesia and various other warmer sea coasts of the world.
Only a few species, however, are found outside Australia.

Calogyne with three species occurs in Australia (2 spp.), and on the coasts
of China (1 sp.).

Selliera with 2 species, occurs in Australia, one species being endemic, while
the other extends to New Zealand and extra-tropical South America.

Willis (1919) gives a larger number of species in three of the above mentioned
genera, viz. Goodenia 100 species, Dampiera 35 species and Verreuxia 3 species,
while the Index Kewensis supplies the following: Goodenia 101 species, Dampiera
48 species, and Verreucia 2 species.

Manifestly, then, Australia is the headquarters of the Goodeniaceae, since
only three of the twelve genera, and these comprising only a few species, occur
outside its borders.

The Goodeniaceae in conjunction with the Cucurbitaceae, Campanulaceae,
Candolleaceae, Calyceraceae, and Compositae form an Order—the Campanales

BY P. BROUGH. ; 473

(Engler and Prantl, 1889)—and it will be interesting to compare the life-history
of one of the species of the Goodeniaceae with the life-histories representative of
other families in the Order.

The vegetative features and general floral structure alike indicate that the
Goddeniaceae are highly specialized, and relatively recent in evolution.

Genus DAMPIERA.

The genus Dampiera has been selected as the type for this investigation
primarily because it is endemic to Australia, but also on account of its wide
distribution throughout the Continent, and its highly specialized floral structure.

Seven of the thirty-four species occur in New South Wales. Two of these
are common in the neighbourhood of Sydney, viz. D. stricta and D. Brownii, the
former providing the material for this research.

Dampiera stricta is typically found growing in the bush in poor sandy soil,
and in distribution ranges from the. coastal districts to the dividing ranges.
Reference to Plate xxxvi shows the habit to be erect, shrubby, and almost
herbaceous. Normally it attains a height of from twelve to eighteen inches.
The stem and leaves are of a dark green colour. The former is distinctly angular,
while the alternate exstipulate leaves vary from elliptical to almost linear in
shape, with the lower ones broader, toothed or entire. The leaves vary from one-
half to one and a half inches in length.

The flowers which are shortly pedunculate, occur mostly in the axils of the
upper leaves, but may be terminal and are either solitary or in clusters. The
number and relative positions of the various floral organs are shown in Text-
figs. 8-11.

The brownish gamosepalous calyx which is partially fused with the ovary
is densely tomentose, and terminates in five tapering lobes. The zygomorphic
corolla is blue with a yellow centre and is about three-quarters of an inch in
length. This corolla is five-lobed and split deeply in the transverse plane in such
a way that two almost free perianth segments occupy an adaxial position while
three sympetalous segments are on the abaxial side. The infolding adjoining
edges of the two adaxial segments form a pouch-like structure. All lobes are
densely hairy except for the membranous margins which are glabrous.

The androecium consists of five stamens which alternate with the perianth
segments, the filaments being inserted singly on the receptacle while the anthers
are connate.

The ovary is bicarpellate, inferior, possesses one loculus, and contains a single
erect anatropous ovule. When the flower is opening the style elongates rapidly,
carries the terminal indusium up through the connate anthers, and curving
slightly near the distal end pushes the indusium into the pouch-like receptacle
formed by the auricles in the line of junction between the two adaxial segments
of the corolla.

Organogeny. }

The four whorls of floral organs—sepals, petals, stamens and carpels—arise
in acropetal succession. The members of any one whorl alternate with the members
of the whorl immediately above or below. This latter fact is shown in Text-figs.
8-11 which represent a flower bud in transverse sections. These sections are
arranged in basipetal sequence.

474 STUDIES IN THE GOODENIACEAE, i,

In Text-fig. 3 at t. is shown the thalamus of a very young flower bud in
longitudinal section. The primordia of the first or sepaline whorl are just
apparent.

Text-fig. 1 shows a young bud in longitudinal section. The young sepals,
petals and primordia of the stamens are easily recognizable, and are quite distinct
from each other.

Text-fig. 1. A median longitudinal section of a very young flower
bud showing young sepals, petals and stamens. s., sepals; p., petals;
st., stamen primordium. x 33.

Text-fig. 2. A median longitudinal section of a flower bud slightly
older than that in Text-fig. 1. In this case the primordia of the carpels’
are apparent. s., sepal; p., petal; st., stamen; c., carpel. x 33.

Text-fig. 3. A median longitudinal section of a still older flower
bud A, also young floral axis B. s., sepal; pl., petal; st., stamen; sp.,
sporogenous cells; sty., style; ov., cavity of ovary; n., nucellus; p.,
primordium of sepal on floral receptacle; t., thalamus. x 33.

A slightly older bud shows the organs just mentioned, but in this case
they have increased in size, and the primordia of the carpels are now apparent,
Text-fig. 2. Even at this early stage the characteristic hairs in the distal region
of the petals are apparent. Text-figs. 4-7 form a series of longitudinal sections
of successively later stages in floral development. In the mature flower the
sepals are fused, but the explanation supplied above indicates clearly that the
primordia of these floral organs arise separately on the thalamus, and that fusion

BY P. BROUGH.

Text-fig. 4. A median longitudinal section of a flower bud slightly
older than that of Text-fig. 3. s., sepal; p., petal; st., stamen; .t.,
microspore tetrads; i., indusium; »., nucellus; f¢., integument. x 25.

Text-figs. 5, 6 and 7. Series of longitudinal sections of flower buds
showing various stages in development. m.s., microspores; interpretation
of other parts as in Text-fig. 4. x 25.

476 STUDIES IN THE GOODENIACEAE, i,

is brought about later by an increase in extent of the various merismatic regions
until they coalesce, and produce the gamosepalous calyx. In the case of the
corolla the primordia also arise separately and continue growing independently
until five distinct lobes are produced (Text-fig. 9). Later the three abaxial segments
fuse (Text-figs. 10 and 11), thereby producing the lower lip of the mature
structure, The two adaxial primordia, however, retain their identity until
maturity is almost attained. Thus, in the fully developed flower, the corolla
consists of a lower lip composed of three fused segments and an upper region,
the two segments of which are distinct from the abaxial lip, and almost from
each other.

Text-figs. 8-11. Series of transverse sections in basipetal sequence
of a flower bud about the same age as that shown in Text-fig. 7. The
relative positions of the various parts, and their gradual fusion is
illustrated. s., sepal; p., petal; st., stamen; sy., style. x 16.

The development of the androecium, however, forms a distinct contrast. In
this case there is no increased merismatic activity in the region of the thalamus
which gives rise to the androecium, and so the primordia of the stamens always
remain distinct from each other. At a later stage the anthers increase in size
with extraordinary rapidity, and the pressure set up by the impinging cells of
the various stamens results in the connate condition seen in the anthers at the
stage of development immediately preceding the final and very rapid elongation
of the style. (Text-figs. 8 and 20).

In the young flower bud (Text-fig. 3) there is no indication of an indusium
or pollen cup. As development proceeds the apex of the style becomes slightly
concave (Text-figs. 4, 6 and 7), owing first of all to the relatively slow growth now
taking place at the organic apex, and secondly to the increased activity of the
cells around the margin of the style apex. This differential growth eventually

BY P. BROUGH. 477

results in the formation of the pollen cup characteristic of the more mature
flower bud (Text-fig. 53).

Again reference to Text-figs. 3-7 shows that during floral development there
are from time to time most striking differences in the relative rates of growth of
the style and stamens. The result of the phenomenon is that at several stages of
floral development the anthers overtop the style apex, while at other intervening
periods the style rises above the stamens. However, before the bud is ready
to open the connate anthers overtop the apex of the style, the extremity of which

Text-fig. 12. Longitudinal section of young anther. av., arche-
sporium; p.l., primary parietal layer; s.l., primary sporogenous layer
cell; c., young carpel. x 266.

Text-fig. 13. Longitudinal section of anther slightly older than
that of Text-fig. 12. w., wall of sporangium; ¢., tapetum; m.c., spore
mother cells. x 266.

Text-fig. 14. Longitudinal section of anther slightly older than that
in Text-fig. 13. ¢., tapetum; m.c., spore mother cells. x 266.
Text-fig. 15. Spore mother cells in synapsis. x 600.

478 STUDIES IN THE GOODENIACEAE, i,

is cup-shaped. The anthers are now mature, and the stimulus exerted by the -
friction of the style in passing up through the mature anthers causes the pollen
sacs to split longitudinally along their inner surface and dehisce introrsely.
Consequently the cup-shaped apex of the style becomes filled to overflowing with
pollen grains. (Plate xxxvii, figs. 3, 4 and 5). Surely it would be hard to find
a more exact or a more wonderful device in pollinating mechanisms. It may also
be observed that only a very small proportion of the microspores are left behind
in the pollen sacs.

Within the stamen whorl, and around the base of the style five nectaries are
situated. One of these organs is shown in detail in Text-fig. 58. Such structures
obviously find their role in connection with insect visitation.

A comparison between the development of the stamens and carpels in point
of time shows that the procedure is normal, the former anticipating the latter
stage by stage until these organs are mature. For example, the archesporial cells
of the microsporangium are differentiated, while the tissue of the megasporangium
is still quite homogeneous and before the integuments make their appearance
(Text-figs. 3 and 12) ; microspore mother cells are produced before the archesporium
of the nucellus is apparent (Text-figs. 13, 14,15); pollen tetrads are segregated by
the time the megaspore mother cell is distinguishable (Text-figs. 4, 26 and 27);
the uninucleate microspore stage is attained, while the megaspore mother cell is
dividing and giving rise to the four megaspores; dehiscence of the microsporangia
and the deposition of the binucleate pollen grains in the cup-shaped indusium of
the elongated style has occurred by the time the mature female gametophyte has
been formed; the 8-nucleate stage of the female gametophyte is not attained until
the flower bud is ready to open.

Although the microspores in the pollen cup are fully developed at this stage,
they do not germinate, the stigma not yet being fully developed or receptive.

The Microsporangium—Development and Structure.

The anther consists of four microsporangia held together by connective tissue
of the filament. The anthers become connate by fusion of the cuticle of contiguous
cells when the stage of development—tetrad stage—shown in Text-fig. 4 is reached.
The fragile nature, and the very restricted amount of tissue controlling this
cohesion are illustrated in Text-fig. 20.

A longitudinal section through the archesporium is shown in Text-fig. 12. The
archesporial cells each divide once by periclinal walls giving rise to the primary
sporogenous layer (s.l.) and the primary parietal layer (p.l.). The latter divides
producing the tapetum on the inside, and a wall layer one cell thick on the
outside (Text-fig. 13).

Later the inner wall layer by differential thickening becomes the fibrous layer,
while the multinucleate tapetal cells become absorbed by the developing micro-
spores. Vestiges of the tapetum are still distinguishable in Text-fig. 20.

The sporogenous cells divide twice or thrice in giving rise to the spore
mother cells.

At the stage when the microspore mother cells are produced, a distinct
tapetum with multinucleate cells, and dense granular cytoplasm is present (Text-
fig. 13). This in turn is encased by the wall of the sporangium which on the
outer side is two cells thick. Most of the spore mother cells, as shown on a
higher scale of magnification in Text-fig. 15, are in the condition known as
synapsis, the nuclei being relatively large, and the chromatin drawn together.

BY P. BROUGH. 479

This phase is quickly followed by that indicated in Text-fig. 16, where short
thick individual chromosomes can be discerned, and where various stages in
reduction division are evident. These cells are found in a longitudinal section
of a single microsporangium, and a Series illustrating the chief phases in reduction
division may be recognized on passing from one end of the section to the other.
This testifies to the rapidity in development of the organ concerned. After the
separation of the chromosomes, daughter nuclei are formed (Text-fig. 17). The

Text-fig. 16. Longitudinal section of microsporangium containing
microspore mother cells with nuclei in various stages of reduction
division. x 600.

Text-fig. 17. Binucleate condition resulting from reduction division
in microspore mother cells. The spindle fibres are still visible in some
cases. x 600.

Text-fig. 18. The four-nucleate stage preceding the formation of the
microspore tetrad. The spindle fibres are still evident. x 600.

Text-fig. 19. Microspore tetrad stage. Only three of the four young
microspores are shown. x 600.

480 STUDIES IN THE GOODENIACEAE, i,

two daughter nuclei in various stages of completeness are evident, the spindle
fibres still persisting. This phase is of very short duration, and is followed with-
out interval by division of the daughter nuclei, so that the simultaneous production
of the four nuclei of the tetrad is effected (Text-fig. 18). The fibres indicating the
existence of the spindle between such nuclei are quite evident, and the view shown
in Text-fig. 18 is taken from another part of the same microsporangium as that
portrayed in Text-fig. 17. This again testifies to the extreme rapidity of the
development of the contents of a microsporangium. The fibres soon disappear,
each nucleus is invested by its share of cytoplasm, and a wall forms round each
young cell, and thus the pollen tetrad stiil enclosed in the common wall of the
mother cell, is evolved (Text-fig. 19). Eventually the pollen grains are set free
by the dissolving of the enclosing wall. The tetrads are produced while the flower-
bud is at the relatively early stage of development shown in Text-fig. 4. Thereafter
the nucleus of each microspore divides (Text-fig. 21, A) and the binucleate
condition of the pollen grain is realized (Text-fig. 21, B). Soon after, but in
many cases before the opening of the flower bud, four equidistant protuberances
are seen to have arisen from the wall of the microspore, thus imparting the
characteristic tetrahedral shape of the mature pollen grain (Text-fig. 21, C).

ea)
Dd
Rg

wy f ORS
ri

t,

SA

iW K(

WF

Text-fig. 20. Transverse section of one anther and part of another
at the stage immediately preceding dehiscence. e., epidermis; f.,
fibrous layer; m., binucleate microspores; ¢t., remains of tapetum;
d., unthickened cells of fibrous layer indicating region of dehiscence;
c., connate cells belonging to two adjacent anthers. x 110.

During microsporogenesis well marked changes fail to be recorded in the
development of the tapetum, and wall of the sporangium. As the stamen
approaches maturity the cells of the inner layer of the wall—which is only two
cells thick—increase in size, and become possessed of a well-marked fibrous
thickening (Text-fig. 20). The cells of the outer or epidermal layer, however,
shrink, especially those on the side of the microsporangium adjoining the style,
and when the time for dehiscence arrives, these epidermal cells have almost
collapsed and form no significant part of the mature structure. The fibrous layer
completely surrounds the microspores except at the line of dehiscence, and is two
cells thick in the region adjacent to the connective. An examination of the fibrous
layer reveals the modus operandi of dehiscence. The partition wall separating
the pairs of microsporangia has already been ruptured, resulting in the formation
of two pollen sacs in each anther. The tapetal cells have meantime been absorbed,

BY P. BROUGH. 481

their substance being used in the nourishment of the developing pollen grains,
and only remnants of the tapetum are now visible. In transverse section the
fibrous layer is seen to be complete except in the region d., where two impinging
cells (one from each microsporangium of the pollen sac) remain unthickened.
Thus a region of weakness extends the whole length of the sac, and when dessica-
tion occurs in the mature sacs a strain is set up in the wall, which lends itself
to ready dehiscence. It will be obvious that were dehiscence to occur while the
anthers still overtop the apex of the style, the microspores shed must drop into
the hollow of the indusium. But at this stage, the elongating style begins to
push the cup-shaped indusium up through the narrow passage between the connate
anthers, and in no case have pollen grains been found in this cup prior to its
passage up through these anthers. Accordingly, the friction set up by the
indusium rubbing against these anthers is the stimulus which determines the
actual time of dehiscence. It is worthy of note that this rapid elongation of the
style terminates in the opening of the bud, and at the time when the microspores
have just reached the binucleate condition. The full significance of this exact
mechanism for abstracting the contents of the pollen sac, and the placing of the
pollen grains in a position favouring dispersal by visiting insects, will be
appreciated later when the problem of pollination is under discussion.

The Male Gametophyte.
The flower of Dampiera stricta is markedly protandrous, and the uninucleate
microspores are produced in the young flower bud (Text-fig. 5). Soon after, and
before the flower opens the nucleus undergoes division (Text-fig. 21, A) and the

Text-fig. 21. Three consecutive stages in development of male
gametophyte. x 600. A. Nucleus of microspore dividing. B. Binucleate
stage. v., vegetative nucleus; g., generative nucleus. C. Tetrahedral
condition of mature microspore with slight protrusion of intine at three
of the four corners.

Text-fig. 22. Two microspores m. and m.,, with pollen tubes p. and
p.,, passing down among receptive papillate cells of stigma. x 266.

binucleate stage is reached (Text-fig. 21, B). Meanwhile the outer wall of the
spore is steadily increasing in thickness until a relatively thick exine is produced.
This thickening is not uniform, however, and in the mature pollen grain four
thin areas are discernible (Text-figs. 21, B, and 21, C). By the time the flower
bud opens the pressure of the protoplasm within the pollen grain has in many
cases increased to such a degree that four distinct protuberances are produced by
the protrusion of the spore wall at the four unthickened regions. This gives the
microspore the tetrahedral shape characteristic of the mature pollen grain.

In order to study the further development of the male gametophyte a number
of mature microspores were removed from the flower and placed in a five per cent.

482 STUDIES IN THE GOODENIACEAE, i,

solution of cane sugar contained in a well-slide. This preparation was then placed -
under the objective of a microscope. After half an hour one of the four pro-
tuberances in each of several microspores under observation was seen to have
increased in size as shown in Text-fig 23, a. Thereafter the young pollen tube
steadily increased in length, and its appearance at intervals of fifteen minutes
was noted and drawn (Plate xxxvii, fig. 6). Thus was obtained the series of

)
ehhh
eshdb odd

Text-fig. 23. Series showing development of pollen tube in a five
per cent. sugar solution. x 225 (approx.).

drawings indicated in Text-fig. 21, a.-j. The last drawing of the series, 21 j.,
. accordingly represents the development attained in 2 hours 45 minutes. It will
be observed that the other three protuberances steadily contract as germination
proceeds, and eventually disappear. The pollen tube therefore develops with great
rapidity, and several such tubes may be detected when a mature style is teased
out under the microscope.

a
Qa fatal 4

panes

pleas fal

Text-fig. 24. Pollen tube within ovary, and passing from style to
miecropyle. w., wall of ovary; o., ovule; m., micropyle; p.t., pollen tube;
e.s., embryo sac. x 16.

Text-fig. 25. Median longitudinal section of ovary showing
anatropous nature of young ovule. i., single massive integument; .i.,
micropyle; m., megasporangium; w., wall of ovary. x 66.

Further development was traced by examining longitudinal sections of the
ovary. In Text-fig. 24, a pollen tube is seen passing from the base of the style to
the micropyle of the ovule.™In this case only a single pollen tube is shown, but
in nearly every case examined several pollen tubes were in evidence. These
either grew along the surface of the ovule or adhered to the inner wall of the

BY P. BROUGH. 483

Ovary until the region of the micropyle was reached. One of the pollen tubes
eventually makes its way through the micropyle and enters the embryo sac in the
region of the egg-apparatus.

The Megasporangium.

The gynoecium is composed of two carpels, the origin and early development
of which have already been referred to. The young carpels arise separately, but
as growth proceeds fusion occurs, and a common style, stigma, and ovary arise.
Growth at the apex of the thalamus is, however, relatively slow, and so the
carpel producing region is gradually overtopped by the stronger growth around
it. The ovary thus becomes inferior, but the style and stigma protrude above the
encircling thalamus. The cavity of the ovary is shown in Text-fig. 3, with a young
nucellus arising therein. So far there is no indication of an integument. A
slightly older stage is shown in Text-fig. 4. Even at this early stage the anatropous
nature of the mature ovule is foreshadowed. A single massive integument has

Text-fig. 26. Median longitudinal section of very young ovule. The
thick integument partially enclosing the megasporangium is seen. i.,
integument; m., megasporangium; m.c., megaspore mother cellwst,
funiculus. x 266.

made its appearance, and partly encloses the nucellus which forms a relatively
small portion of the ovule. Text-fig. 26 illustrates the structure of the very
young ovule, at a stage slightly younger than that just referred to. The
funiculus, integument and megasporangium are apparent, the last named consisting
of an axial row of cells surrounded by a jacket one-cell thick. The megasporangium
is therefore of the reduced type characteristic of the more highly developed
Angiosperm families. As development proceeds the integument soon outstrips the
nucellus, which becomes completely enclosed except for the micropylar passage
at the distal end. Various stages of this envelopment are illustrated in Text-
figs. 4, 5, 6, 7, 25, 26 and 29. The long micropyle leading from the apex of the
nucellus is shown in Text-figs. 25 and 29. It is to be noted that thus early the
cells of the integument bordering on that region of the micropyle adjacent to the
megasporangium are densely cytoplasmic, and clearly in a state of high nutrition
(Text-fig. 29). Later on, the functional megaspore and embryo sac grow forward
into the micropyle, and eventually attain a position just beyond this nutritive
jacket, which functions as a tapetum, a feature not uncommon in representatives
of the more highly developed Angiosperms.

484 STUDIES IN THE GOODENIACEAE, i,

In dealing with this structure Coulter and Chamberlain (1915) point out that -
it is usually derived from the integument, but arises from the nucellus in Armeria.
They observe: “This jacket has been definitely observed as conspicuous in Helosis
(Chodat and Bernard, 1900), Sium, many Scrophulariaceae (Balicka-[wanowska,
1899), Campanula (Barnes, 1885), Stylidaceae (Burns, 1900) and certain

Text-fig. 27. Median longitudinal section of megasporangium. The
large cell terminating the axial row of the megasporangium is the
megaspore mother cell, m.c.. x 450.

Compositae, and by Billings (1901) in numerous Sympetalous forms, among the
most conspicuous being Lobelia. Primulaceae (except Leptosiphon), Linum,
Forsythia, Amsonia, Menyanthes, Polemoniaceae, Myoporum, Globularia, Scaevola,
Calendula, etc.”.

From this list it is evident that a nutritive jacket around the embryo sac is of
frequent occurrence among the Sympetalae (especially amongst the more highly
developed families) but is also known amongst the Archichlamydeae though to
a much less extent. ;

Just when the anatropous nature of the ovule has become apparent, it is seen
that the distal cell of the axial row of the nucellus is large relative to the others,
and has a nucleus in proportion. This cell is clearly a megaspore mother cell,
and it continues to enlarge until of the dimensions shown in Text-figs. 26 and 27.
The nucleus is in synapsis, but examination of numerous longitudinal sections of
buds about this stage of development, failed to demonstrate actual reduction
division. However, a subsequent stage is indicated in Text-fig. 28 where the four
nuclei of the megaspore tetrad are visible. These are not separated by cell walls.
A further stage in development is shown in Text-fig. 30 where the two nuclei
towards the distal end are quite evident, but the third is undergoing disintegration.
The reason for this is seen when the fourth, innermost, or chalazal nucleus is
examined. It easily exceeds all the others in dimensions and is clearly the
functioning megaspore. Its growth has resulted in the partial disintegration of
the adjacent nucleus which is seen in a matrix of deeply stained matter. The
steady increase in size of the functional megaspore results in the disorganization
of the other three megaspores as illustrated in Text-fig. 31, where only a remnant
of the distal nucleus is discernible, the other two non-functional megaspores having
completely disappeared. Their former position is occupied by a dark coloured
mass which stains very deeply with Haidenhain’s iron alum and Fleming’s triple
stains respectively. Eventually this mass—which evidently is used in the

BY P. BROUGI. ; 485

nutrition of the functional megaspore—completely disappears, and the young
embryo sac with its large nucleus is evolved (Text-fig. 32).

The Female Gametophyte.
Meanwhile the nutritive jacket is becoming more and more evident, extending
well down the micropyle, and also completely surrounding the megasporangium.

Text-fig. 28. Median longitudinal section of slightly older ovule
showing 4 megaspores not separated by walls. x 266.

Text-fig. 29. Longitudinal section through micropyle showing
nutritive jacket, n.j., formed by innermost cells of integument. The
megasporangium, m., is not quite in median section. x 266.

Text-fig. 30. Median longitudinal section of megasporangium
showing enlarged functional megaspore, m., and the adjoining
megaspore, m.,, disintegrating. x 333.

Text-fig. 31. Median longitudinal section of megasporangium
showing functional megaspore, m. The disintegration of the other three
megaspores is almost complete. x 333.

486 STUDIES IN THE GOODENIACEAE, 1,

As the functional megaspore increases in size, the single layer of cells forming the
wall of the megasporangium is steadily encroached upon and absorbed (Text-
fig. 32), so that by the time the young embryo sac has been formed, the wall
cells have practically disappeared. The nucleus of the surviving megaspore now
divides, producing the first two nuclei of the female gametophyte (Text-fig. 33).

Text-fig. 32. Longitudinal section of ovule showing functional
megaspore, m#., and remains of nucellar tissue now almost absorbed by
developing embryo sac. x 300.

Text-fig. 33. Portion of median longitudinal section of an ovule
showing binucleate condition of embryo sac. wn.j., nutritive jacket.

x 300.
Text-fig. 34. Binucleate condition of embryo sac slightly older than

that figured in previous drawing. x 310.

The embryo sac and nuclei enlarge rapidly, the latter being gradually separated
(Text-fig. 34), and eventually occupying opposite ends of the sac. Each nucleus
divides again producing the four-nucleate stage illustrated in Text-fig. 35. A
subsequent division of each of these nuclei results in the eight-nucleate stage of the
mature gametophyte. Text-figs. 36 and 37 respectively show the micropylar portions
of the two embryo sacs. In each case the egg apparatus is depicted, while in the
former the endosperm nucleus is represented.

Text-fig. 38 shows three successive sections of a mature and enlarged embryo
sac. In B a single synergid and the egg are apparent; in A a second synergid
and the micropylar polar nucleus are evident; while in C the antipodal polar nucleus
and the three antipodal cells are found. The synergids are pyriform, their

5
(

BY P. BROUGH. 48

tapering ends invade the micropyle to some slight extent, and each possesses a
nucleus and a large vacuole. The oosphere is apparent at the lower extremities
of the synergids. The tapetum is figured on one side of the embryo sac only. This
nutritive jacket is now at the height of its development, and its dense deeply

Text-fig. 35. The four-nucleate condition of the embryo. sac.
m., micropylar region of sac; «a., antipodal region of sac. x 266.
Text-figs. 36 and 37. Micropylar region of two embryo sacs.
S.,, Synergids; o., oosphere; f., primary endosperm nucleus. x 266.

stained cytoplasm, coupled with the regular arrangement of its constituent cells
brings it out in bold relief in the sections. It will be seen that the micropylar
region of the sac has grown slightly beyond the tapetal cells of this region. A
slightly older stage of the sac is seen in Text-fig. 39, A. Therein the egg, the fusing
polar nuclei, and two of the antipodal cells are delineated. Part of the pollen tube
is visible, and what are interpreted as the two male nuclei—one in contact with
the egg and another with the fusing polar nuclei—are indicated.

It must be admitted, however, that these preparations were not distinct
enough to diagnose the male nuclei with absolute certainty nor to place the
behaviour of the male nuclei beyond the possibility of doubt. The view of what
is regarded as the triple fusion nucleus, as obtained under the oil-immersion lens,
is shown in Text-fig. 38, B.

The behaviour. of the polar nuclei, however, presented no difficulties, as their
history could be traced with considerable ease. Their gradual fusion is illustrated
in Text-figs. 40-48, but in none of the cases illustrated is the second male nucleus
in evidence.

488

STUDIES 1N THE GOODENTACEAE, 1,

Text-fig. 38. Text-figs. 38A, 38B and 38C show views seen in
three successive sections of embryo sac. s.,, synergid; s.,, synergid;
o., oosphere; f.,, micropylar fusion nucleus; f.,, antipodal fusion nucleus ;

a., a., and a.,, antipodal cells; n.j., nutritive jacket shown on one side
of embryo sac only. x 266.

Text-fig. 39. A. Another embryo sac. o., oosphere; e.n., endosperm
nucleus; a., antipodal cells; p., pollen tube. x 266. B. Fusion of polar
nuclei, p., and p.,, with male nucleus, ». x 600.

BY P. BROUGH. 489

The chalazal region of the embryo sac is occupied by three antipodal cells.
These are well developed and vary in shape and size, the one occupying the lower
extremity of the sac being somewhat elongated, while the other two are almost

Text-figs. 40-43. Polar nuclei in various stages of fusion. x 300.

isodiametric. These cells remain active until after fertilization, evidently
functioning in passing nourishment from the chalaza of the ovule into the
developing sac, but unlike the antipodals found in many of the higher Sympetalae,
e.g. Aster novae-anglicae (Chamberlain, J. C., 1895), Sherardia arvensis (Lloyd,
F. E., 1902), do not become unduly enlarged, nor do they encroach to any extent
on the chalazal region as haustorial invaders.

The endosperm nucleus divides before that of the fertilized egg or oospore
giving rise to free endosperm nuclei as demonstrated in Text-fig. 44. The embryo
sac is long and narrow at this stage, so that comparatively few free endosperm
nuclei are formed before wall formation is initiated. This is accompanied by rapid
expansion of the sac. The nutritive jacket still persists unimpaired.

Emobryogeny.

When the oospore divides and commences to produce the embryo, a large
mass of endosperm tissue awaits invasion. The details of embryogeny are
illustrated in a series of drawings (Text-figs. 45-52).

Actual division of the oospore has not been demonstrated, but Text-fig. 45
shows the stage immediately subsequent. Examination reveals a pro-embryo of
two cells—a basal or embryo cell proper, and a suspensor cell. The nucleus of
the latter is in mitosis.

The pro-embryo invades the endosperm tissue, the cells of which are being
absorbed. The cells of the young embryo evidently secrete an enzyme which
gradually brings the contingent endosperm cells into solution, preparatory to
absorption. The digestive action of the embryo is indicated by the corroded
appearance of the adjacent endosperm cells. This feature is apparent in Text-
figs. 45, 48 and 51.

The cells of the nutritive jacket still preserve their form, but are now
distinctly vacuolate. Text-figs. 45, 46 and 48 show the gradual production of the
suspensor which pushes the embryo cell well down amid the endosperm cells.

490

STUDIES IN THE GOODENIACEAE, 1,

Text-fig. 44. A longitudinal seetion through an embryo sac after
fertilization. s., synergid; o., oospore; f.n., free endosperm nuclei.
x 266.

Text-fig. 45. Very early stage in embryology soon after division
of oospore. e¢., embryonal cell; s., first cell of suspensor with nucleus
undergoing division; n.j., nutritive jacket; c., corroded cells of endosperm
tissue. x 600.

BY P. BROUGH. , 491

Text-fig. 46. <A slightly older stage of embryo. The suspensor has
elongated. e., embryonal cell; s., suspensor; ev., endosperm; 11.j..
* nutritive jacket. x 266.
Text-fig. 47. A further stage in development of embryo. The
embryonal cell has divided and approached the octant stage, 0. x 600.

Text-figs. 48-50 show further stages in embryogeny. x 266.

Text-fig. 51. Embryo, e., invading the endosperm, en.; 12.j., nutritive
jacket around sac. x 150.

Text-fig. 52. Late stage in embryogeny. The dermatogen, periblem
and plerome are differentiated. x 266.

492 STUDIES IN THE GOODENIACEAF, 1,

Finally the terminal cell divides so that the quadrant stage, and later the octant .
stage, are arrived at (Text-fig. 47).

Further cell division follows the octant stage, as indicated in Text-figs. 49-52,
and eventually the relatively mature condition depicted in Text-fig. 52 is arrived
at. Therein the dermatogen is well defined, and the differentiation into periblem
and plerome suggested. Beyond this stage the development of the embryo was
not traced.

Pollination.

The problem of pollination and the mechanisms incidental thereto in the
Goodeniaceae have attracted widespread attention. Several investigators—notably
Brown, R. (1818, 1866), Mueller, H. (1883), Haviland, H. (1884, 1885), Haviland,
F. E. (1914), and Hamilton, A. G. (1885, 1894), have placed the results of their

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Text-fig. 53. Longitudinal section of apex of style. The young
stigma, s., is growing up from the base of the pollen cup, and forcing
out the microspores through the apical pore, p. x 73.

Text-fig. 54. Another view of same showing conducting tissue of
style. The portion of the cup r. has been slightly displaced in cutting
the section. x 73.

Text-fig. 55. Part of stigma of previous figure shown under a
higher power of magnification in order to illustrate nature of component
cells. x 133. :

Text-fig. 56. Longitudinal section of an older stage of style showing
stigma occupying centre of pollen cup. The bi-lobed nature of stigma,
and the central papillate cells, p., are evident. x 33.

BY P. BROUGH. 493

investigations on record in the case of certain species of Goodenia. The
conclusions arrived at are so varied, however, that a careful study of the method
of pollination in other genera of the family seems advisable.

In the case of Dampiera stricta the method of dehiscence of the anthers, and
the depositing of the pollen grains within the hollow indusium have already been
described. The cup is filled to overflowing, the spores forming a pyramidal mass.
Thereafter the indusium closes owing to the contraction of the margins.

At the time when the style finally elongates, and the flower bud is on the point
of opening, the indusium is forcibly thrust into a chamber formed by the auricles
in the line of junction between the two posterior petals. The two flap-like
appendages aid in the complete enclosing of the indusium, which is orientated so
as to face the anterior side of the flower. At no period then, has the pollen been
exposed, although part of the bent style is visible, outside the corolla between the
two posterior petals. By this time the anthers have withered, and are of no
further significance. The rudimentary stigma at the base of the closed indusium
now commences to grow rapidly (Text-figs. 53 and 54) and gradually pushes the
pollen in front of it, and out through the narrow aperture now present at the top
of the fringed pollen cup (Plate xxxvii, figs. 3, 4 and 5). The pollen thus ejected
either lodges on the outer sloping sides of the indusium, as indicated in the
photographs just referred to, or is deposited in the chamber in which the indusium
is imprisoned. Obviously such pollen grains cannot regain their former position
resting on the apex of the stigma. Eventually the steadily expanding and rounded
stigma protrudes above the rim of the inner cup of the indusium (Text-fig. 56)
and occupies the cavity formerly filled by pollen grains. A few microspores may
remain lodged between the lower portion of the stigma and the inner wall of the
indusium, but such are ineffective from the point of view of pollination. The
stigma finally becomes slightly two-lobed, and the cells within and around the
depression so formed become papillate, and eventually receptive in relation to the
pollen grains (Text-figs. 56, 57 and 22). Furthermore, the sub-epidermal cells
immediately within this cleft, and also the tissue leading therefrom, and down
through the centre of the style are of a loose, thin-walled nature. Such cells
have all the characters of a typical conducting tissue for the nutrition of the
advancing pollen tube. Pollen tubes in considerable number were found when the
central tissue of the style was dissected out, and examined under a microscope.

In no case were microspores seen to germinate while still within the indusium,
although at this stage they are in the binucleate condition, and in many cases
show the slight protrusion of the intine through the four weak areas in the exine
characteristic of the mature microspore. Judging from the various stigmas
examined, at different stages of growth up to and including the emergence of the
stigma from the indusium, it seems clear that the pollen grains do not germinate
within their own indusium.

Evidently, then, the stigma does not become receptive until the pollen grains
from the same flower have been ejected from the pollen cup.

Now, five well developed nectaries are found at the base of and within the
whorl of stamens. These are obviously related to insect visitation, and it is well
known that bees are frequent visitors to the flower in question. That insects are
instrumental in the transference of pollen is proved by the fact that in one case
foreign pollen was actually found on a stigma examined. The contrast which
this pollen (which was round and spiny) made with the characteristic microspores
of Dampiera, rendered this point peculiarly easy of elucidation. The foreign

494 STUDIES IN TILE GOODENIACEAE, i,

pollen had not germinated. From what has already been demonstrated in regard .
to the rate of growth of the pollen-tube (Text-fig. 23), it follows that fertilization
may occur within a few hours of the pollen being deposited on the ripe stigma.
An insect entering the flower pushes the two adaxial perianth segments apart,
and, at the same time releases the pollen cup from its chamber. Pollen thus falls
on the visitor, and such action will continue until all the pollen has been removed,

Text-fig. 57. Papillate cells of stigma under higher power of
magnification. x 266.

Text-fig. 58. Longitudinal section of nectary, n., situated at base of
style, s., 2nd within filament of stamen, f. x 150.

a consummation which may well be effected before the stigma becomes receptive.
Pollen from the pouch is also removed at this stage. It was also observed that a
considerable proportion (about 20% or more) of the pollen grains never reach
maturity. Thus when the stigma becomes receptive the pollen grains have either
been removed by insects or have been pushed aside, and so subsequent insects
will deposit pollen from their backs or heads on the surface of the receptive stigma.

It must be conceded then, that pollen is actually transferred by insects from
one flower of Dampiera stricta to another flower on the same or on another plant,
and so cross pollination is effected.

It is not impossible, however, that self pollination may occur in the event
of the flower not being visited by the insects, although the writer is of the
opinion that the available evidence is against the view that self-fertilization does
actually occur. The point. can be settled only by taking means to prevent the
access of foreign pollen, and then determining whether embryos have formed in
the ovaries concerned. The writer hopes to elucidate this point in the near future.

An experiment of this nature has already been carried out in the case of

BY P. BROUGH. 495

Goodenia cycloptera by Haviland, F. E. (1914), who found that embryos were not
formed.

A survey of the facts laid bare by this investigation of various phases in the
life-history of Dampiera stricta reveals few features differing markedly from those
characteristic of the various forms comprising the higher Sympetalae. The details
of organogeny, microsporogenesis, megasporogenesis and embryogeny conform to
type, and indicate that the Goodeniaceae find their natural position in the order
Campanales. The genus Dampiera despite its being endemic to Australia does not
show any signal aberrant features, such as was laid bare by an examination of
another great Australian family, namely the Epacridaceae, a member of which,
Styphelia longifolia (Brough, 1923, 1924), showed unique characters more especially
in megasporogenesis.

Reverting to Dampiera it is interesting to observe the syngenesious anthers—
a feature so characteristic of the Campanales. The piston-like action of the style
is also typical, but Dampiera with its protandry, pollen cup, delicate anther
dehiscence, method of dispersing the pollen, and the very late exposure of
receptive stigmatic cells is perhaps the most outstanding in an Order noted for
its highly developed pollinating mechanisms. The genus would seem to present
an epitome of the many peculiarities portrayed by the various members of the
most recent and specialized of all the Orders of flowering plants.

On looking for genetic relationships between the Goodeniaceae and the other
members of the Order Campanales certain striking features already noted in
Dampiera are seen to be reflected in the tribe Lobelioideae of the Campanulaceae.
In this tribe one finds the herbaceous habit, the zygomorphic flowers, the blue
corolla often split to the base, the syngenesious anthers, the piston-action of the
style, and the nutritive jacket around the embryo sac. The genus Lobelia is
represented by eighteen species in Australia (Bentham and Hooker), and the
writer has come to the conclusion that all the available evidence goes to show
that the genus Dampiera—if not the Goodeniaceae as a whole—has been derived
from the Lobelioideae.

Summary.

Organogeny. The primordia of the various sets of floral organs arise separately
and in acropetal succession. The five young sepals eventually fuse, and form a
gamosepalous calyx joined to the ovary. The five petal primordia maintain their
identity until an advanced stage of bud development has been attained, when the
three abaxial segments fuse to form the lower lip, while the two remaining adaxial
segments form the upper lip which is split almost to the base. The two lips are
separate throughout. The stamens retain their identity until about the microspore
mother cell stage, when the syngenesious condition arises by fusion of contiguous
cuticles of the anthers.

The two young carpels fuse to form a common style and inferior ovary.

The characteristic pollen cup arises during late bud development by the
relatively slow rate of growth at the organic apex of the style, and the increased
merismatic activity in the marginal region.

Microsporangium. The archesporial cells divide by periclinal walls giving
rise to a primary parietal layer and a primary sporogenous layer. The cells of the
former divide once thereby producing a wall layer and the tapetum. The cells
of the primary sporogenous layer divide two or three times before attaining the
spore mother cell stage.

K

496 STUDIES IN THE GOODENIACEAE, 1,

The spore mother cells undergo reduction division and then ordinary somatic -
division, whereby simultaneous pollen tetrads are produced. The great rapidity
in growth is attested by the wide range in development depicted in the longitudinal
section of a single sporangium. In the mature pollen sac the spores are
binucleate, each spore wall bears four equidistant thin areas, the hypodermal cells
constitute the fibrous layer, and dehiscence is effected by longitudinal splitting
along two median vertical contingent rows of unthickened cells of the hypodermal
layer. The mature stamens are short, and are never exposed outside the flower.

The male gametophyte. The mature tetrahedral pollen grain contains a
generative and a vegetative nucleus. During germination a single pollen tube
grows out through one of the four unthickened areas of the mature microspore.
Evidence testifying to the rapidity in growth of the pollen tube was obtained by
careful observation of its development in a five per cent. sugar solution. Pollen
tubes were traced from the receptive cells of the stigma, throughout the style,
and in the ovary from the base of the style to the micropyle. The tube grows
through the micropyle, and enters the embryo sac in the region of the egg
apparatus.

The megasporangium. A solitary nucellus arises within the ovary. A single
thick integument gradually encloses the young megasporangium, which consists
of an axial row of cells surrounded by a jacket one cell thick. The anatropous
nature of the ovule is early foreshadowed. The megaspore mother cell is formed
at the micropylar end of the axial row of the megasporangium. The mother cell
gives rise to a linear tetrad of megaspores, the innermost of which is the
functional megaspore; this develops rapidly and absorbs the other three
megaspores. The wall cells of the megasporangium are also absorbed. Meantime
the cells of the integument lining the micropyle enlarge, become densely cyto-
plasmic, and eventually form a very definite and noteworthy nutritive jacket
around the embryo sac.

The female gametophyte. The functional megaspore increases in size, and
steadily invades the micropyle. During this development, the bi-nucleate, four-
nucleate, and eight-nucleate stages of the female gametophyte are attained. The
egg apparatus, polar nuclei, and antipodal cells are normal in structure and
polarity. These are described in detail.

Fertilization. The pollen tube enters the embryo sac. One male nucleus
fertilizes the oosphere, while the other joins the partially fused polar nuclei.

Endosperm formation. The endosperm nucleus divides before that of the
oospore—free endosperm nuciei being formed. Wall formation, accompanied
by rapid growth of the embryo sac then supervenes. A massive endosperm is
formed.

Embryogeny. The oospore commences development soon after wall formation
in the endosperm has been initiated. A long suspensor is formed which pushes
the terminal cell well down into the soft endosperm tissue. The embryonal cell
attains the quadrant and octant stages respectively. Later periclinal walls cut
off the dermatogen, and in the final stages examined, the periblem and plerome
had been differentiated.

Pollination. Many investigators, for example R. Brown, H. Mueller, E.
Haviland, F. E. Haviland, and A. G. Hamilton, have carried out investigations
regarding the methods of pollination in members of the Goodeniaceae. Conflicting
views have been expressed. In the case of Dampiera stricta the elongating style
carries the pollen cup up through the syngenesious mature anthers causing

BY P. BROUGH. 497

introrse dehiscence of the microsporangia, and the filling of the cup with pollen
grains. The style further elongates and thrusts the cup into a pouch in the line
of junction of the two adaxial segments of the corolla. The cup is orientated so
as to face the centre of the corolla. The stigma then arises from the base of
the pollen cup, and very gradually forces the microspores out through the pore
at the apex. Such microspores are then in a suitable position for removal on
the backs of visiting insects. Eventually the fully developed and slightly bi-lobed
stigma occupies the whole of the interior of the cup. Then, and not till then,
do receptive cells appear in the apical region of the stigma, which is now ready
to receive pollen from insect visitors coming from less mature flowers.

A very exact mechanism to ensure cross pollination is thus demonstrated. It
would seem, however, that self pollination is possible in the absence of insect
visitors.

The evidence derived from this investigation strongly supports the view that
the genus Dampiera is derived from the Lobelioideae.

List of References.

BALIKA-IWANOWSKA, G. P., 1899.—Contribution & l’étude du sac embryonnaire chez
certaines Gamopetals. Flora, 86, pp. 47-71, pls. 3-10.
BarRNEs, C. R., 1885.—The Process of Fertilization in Campanula americana. Bot.
Gazette 10, pp. 349-354, pl. 10.
BENTHAM AND HooKeEr, 1869.—Flora Australiensis.
Biuuines, F. H., 1901.—Beitr’ige zur Kenntniss der Sammenentwicklung. Flora 88,
pp. 253-318.
BrouGH, P., 1923.—Preliminary note on the Embryo Sac of Styphelia longifolia. Proc.
Linn. Soc. N.S.W., xlviii, pp. 674-680.
, 1924.—Studies in the Epacridaceae. i. The Life-history of Styphelia longifolia.
Proc. Linn. Soc. N.S.W., xlix, pp. 162-178.
Brown, R., 1818.—Queries respecting the Indusium of Brunonia and Goodeniaceae.
Trans. Linn. Soc., xii, p. 134.
, 1866.—Miscellaneous botanical notes, Vol. i, p. 31.
CHAMBERLAIN, C. J., 1895.—The Embryo-sac of Aster novae-anglicae. Bot. Gazette 20,
pp. 205-212, pls. 15-16.
CHODAT, R., AND BERNARD, C., 1900.—Sur le sac embryonnaire de lVHelosis guayanensis.
Jour. Botanique 14, pp. 72-79, pls. 1-2.
CouLTerR, J. M., AND CHAMBERLAIN, C. J., 1915.—Morphology of Angiosperms, p. 1038.
ENGLER AND PRANTL, 1889.—Die Naturlichen Pflanzenfamilien.
HAMILTON, A. G., 1885.—On the Fertilization of Goodenia hederacea. Proc. LINN. Soc.
N.S.W., x, pp. 157-160.
, 1894.—Notes on the Methods of Fertilization of the Goodeniaceae. Proc. LINN.
Soc. N.S.W., xix, p. 201-212.
HAVILAND, E., 1884.—Occasional notes on Plants indigenous in the immediate Neighbour-
hood of Sydney. Proc. LINN. Soc. N.S.W., ix, pp. 449-452.
, 1885.—Some remarks on the Fertilization of the Genus Goodenia. Proc, LINN.
Soc. N.S.W., xX, pp. 237-240.
HAVILAND, F. E., 1914.—The Pollination of Goodenia cycloptera. Proc. LINN. Soc.
N.S.W., xxxix, pp. 851-854.
Luoyrp, F. E., 1902.—The Comparative Embryology of the Rubiaceae. Mem. Torr. Bot.
Club, 8, pp. 27-112.
MUELLER, H., 1883.—The Fertilization of Flowers.
Wiis, J. C., 1919.—Flowering Plants and Ferns.
Index Kewensis.

EXPLANATION OF PLATES XXXVI-XXXVII.
Plate xxxvi.

1. General habit of Dampiera stricta; about one-third natural size
2. Flowers of D. stricta; about one and a half times natural size.

498 STUDIES IN THE GOODENIACEAE, 1.

Plate xxxvii.
Style supporting pollen cup with narrow orifice through which microspores are being
gradually ejected.
4. A style slightly more advanced than that of previous figure. The pollen cup is now
at right angles to main style. The gradual ejection of microspores from the
pollen cup is illustrated.

5. View showing the short stamens arranged around the style. The anthers have
separated, and dehiscence has occurred.

(Je)

6. Microspores germinating. x 100 (approx.).

PLACENTATION AND OTHER PHENOMENA IN THE SCINCID LIZARD
LYGOSOMA (HINULIA) QUOYI.

By H. CriarreE WEEKES, B.Sc., Linnean Macleay Fellow of the
Society in Zoology.

(From the Department of Zoology, University of Sydney.)

(Plates’ xxxviii-xl and twenty-three Text-figures. )
[Read 26th October, 1927.]

Contents.

IT. Introduction.
II. Material and Methods.
III. Description of Material.
1. Period of ‘‘pro-oestrus”’.
2. Period of ovulation and fertilization.
3. Period of placental activity.
Stage A. Early development of the placentae.
Stage B. The stage of placental maturity.
Stage C. The placentae immediately prior to the birth of the foetus.

4. Period after birth.
IV. Comparison with the lizards, Chalcides tridactylus, Lygosoma (Liolepisma),
entrecasteauaxi and Tiliqua scincoides and with the marsupial, Perameles.
V. Theoretical Considerations.
VI. Summary and Conelusions.

I. INTRODUCTION.

In a “Note on Reproductive Phenomena in some Lizards’ communicated to
this Society in May, 1927, the placentation of the Scincid lizard Lygosoma
(Hinulia) quoyi was described briefly and compared and contrasted with that
of the Scincid lizards Chalcides tridactylus, Lygosoma (Liolepisma) entrecasteauxi
and Tiliqua scincoides and with that of the marsupial Perameles. It is proposed
to describe the placentation of L. quoyi and to discuss its relationship with that of
the three above-mentioned Scincid lizards and of the marsupial Perameles in more
detail in this communication.

In a previous paper (Harrison and Weekes, 1925) in which the occurrence of
the placentation of the Scincid lizard L. entrecasteauxri was described, L. quoyi
was mentioned, specimens of both having been collected at Barrington Tops during
January and February, 1925, by the members of a party from the University of
Sydney, under the leadership of Professor Harrison. In this paper (p. 471) it was
stated that examination of pregnant females of Trachysaurus rugosus, T. scincoides,
L. quoyi, Egernia striolata and EH. whitei revealed “highly vascularized external
allantoic and uterine walls with their respective circulations in close apposition
but no marked placental area such as is described below” (for L. entrecasteauw-ri).
A study of the condition in L. quoyi has shown that there is a very definite type
of allantoplacenta present, although there is not the same degree of external
differentiation of the uterine placental area. A further study has shown that a

L

500 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

somewhat similar placental condition occurs in #. striolata and H. whitei and
probably also in 7. rugosus.

Definite placentation among lizards has now been recorded in Chalcides
tridactylus and C. ocellatus by Giacomini (1898) and (1906) respectively; in
Tiliqua scincoides by Flynn (1923); in Lygosoma (Liolepisma) entrecasteauxi by
Harrison and Weekes (1925); in Lygosoma (Hinulia) quoyi, Egernia striolata and
E. whitei by Weekes (1927); and omphaloplacentation in 7’. scincoides by Weekes
(1927). In addition Haacke (1885) mentions certain relations between the wall
of the uterus and that of the yolk-sac in Trachysaurus rugosus. It is probable
that a further investigation of the viviparous reptiles will reveal BEY interesting
phenomena with regard to placentation.

I wish to thank Professor L. Harrison, in whose department this investization
was carried out, for the personal interest he has taken in the work and for the
help he has given; Dr. I. M. Mackerras for specimens of L. quoyi collected at
Barrington Tops; Mr. J. Hopson for collecting and forwarding material; Miss L.
Wood, Mr. W. Graham, Mr. G. Vance and Miss B. White for help in collecting
material; the Department of Zoology, University of Sydney, for grants covering
the cost of collecting.

This investigation was begun under a Science Research Scholarship of the
University of Sydney, and continued after my appointment as a Linnean Macleay
Fellow of this Society.

II. MATERIAL AND METHODS.

Specimens of L. quoyi containing young were collected from Barrington Tops,
150 miles north of Sydney, at a height of 4,500-5,000 feet, during January, February
and December, 1925; from Mount Kosciusko, 300 miles south of Sydney, at a height
of approximately 5,000 feet, during November and December, 1925; from the Biue
Mountains, 70 miles west of Sydney, at a height of approximately 3,000 feet, during
January and February, 1927; from Sydney, at sea level, during November and
December, 1925, and September, October, November, December, 1926, and January,
1927; from Kiama, on the Coast, 70 miles south of Sydney, ayes January,
October and November, 1926.

The study of L. quoyi is interesting apart from the factor of plaventations The
specimens collected from Barrington Tops and Mount Kosciusko, at a height of
approximately 5,000 feet, differ from those collected at sea level, although each
type has been identified as Lygosoma (Hinulia) quoyi. Those collected at
Barrington Tops and Mount Kosciusko have an average length of five inches and
the pregnant females contain from three to five young; those from the coast are
much larger measuring from ten to thirteen inches and the pregnant females
contain from five to nine young. The difference in scale marking between the
two types of lizard is slight, since members of the same type vary among them-
selves, and the placentae in both are identical. The difference in the number of
young carried during the gestation period seems to depend on the size of the
parent lizard, although this conclusion may not be correct, since the increase in
the size of the adult is naturally followed by a corresponding increase in the size
of the embryo, those of the mountain type measuring approximately 3 cm. at the
time of birth, and those of the coastal type measuring approximately 5 cm. Those
females collected at the Blue Mountains, at 3,000 feet, were intermediate in size
between the two above described types measuring on an average eight inches in
length and containing frqm three to five young.

BY H. CLAIRE WEEKES. 501

The reason for the difference in the size of the mountain and coastal types
of lizard is not apparent, but in January, 1927, an intermediate type was collected
at a height of 3,000 feet indicating that altitude may be the determining factor,
and that the change from the small to the large type of lizard may be gradual as
the altitude decreases.

L. quoyi is essentially a water loving animal and always lives near a creek or
water hole. The flat marshy land at Barrington Tops and Mount Kosciusko with
its network of small watercourses is infested with them. They live in holes in
the banks of the watercourses, and at the coast they live in similar holes in the
banks of coastal streams.

As mentioned, pregnant females were collected from Mount Kosciusko during
November and December, 1925, and from Barrington Tops during January and
February, 1925. They were first collected from Mount Kosciusko on the !4th
November and the contained embryos are estimated to be one week old, hence
fertilization occurred during the first week in November. The females collected
from Barrington Tops during the first week in February contained embryos which
were within a few days of hatching, but since it is at present unknown to the
writer whether the times of the fertilization of the ova of the lizards at Mount
Kosciusko and Barrington Tops correspond, the exact length of the gestation period
of the mountain type is unknown.

Females of the coastal type collected at Sydney on 10th September and 7th
October, 1926, had the ova still within the ovaries. On the 18th October females
were collected which had the ova in early stages of segmentation within the
oviducts, so that fertilization in these lizards occurred in the middle of October,
which is two weeks earlier than at Mount Kosciusko. Lizards were collected
from the same locality at Sydney, from the time of the fertilization of their ova
until the time of the birth of the young, which was the second week in January.
Hence the development of L. quoyi at sea level covers a period of approximately
three months. Females were kept alive until the young were born and thus
post partum stages were obtained.

Summarizing the above, specimens of L. quoyi were collected at a height of
4,500-5,000 feet which contained embryos in stages of development ranging from
the first week after fertilization until the time of hatching; specimens were
collected at sea level at all stages ranging from the condition of non-pregnancy to
the condition after birth.

All material was fixed in Bles’ Solution (90 parts of 70% alcohol, 7 parts of
5% formol, and 3 parts glacial acetic acid). The ventral body wall of each female
was cut longitudinally to expose the oviducts, the female with young in situ then
being immersed in the fixative. It is not advisable to leave the material in ‘the
fixative indefinitely as Bles’ Solution hardens yolk.

As most of the material contained much yolk it was found difficult at first
to get satisfactory results when infiltrating without using the method of double
embedding in celloidin and paraffin, a method which has many disadvantages when
dealing with yolky material. Embedding in paraffin alone was successful when
the following precautions were taken: (a) to secure a thorough dehydration of
the material, the latter, if bulky, being passed through many changes of absolute
alcohol over a period of at least two days; (b) to use pure clearing agents,
preferably xylol or cedar wood oil. As an inferior quality of xylol is usually
sold which gives a white precipitate when mixed with alcohol, it is advisable
to use cedar wood oil. However, if the xylol will mix with 70% alcohol without

502 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

giving a permanent precipitate, it will give better results than the cedar wood
oil. Cedar wood oil which is sold as pure often contains water, but it was found
that by mixing the oil with a fair quantity of anhydrous copper sulphate, shaking
well and allowing to stand for twenty-four hours and then filtering, the water and
other impurities were removed and the oil made perfect for use; (c) to infiltrate
gradually, leaving the cleared material at room temperature in a solution of the
clearing agent saturated with paraffin for about twenty-four hours, then leaving
it another twenty-four hours at about 30° C., more paraffin having been added, and
finally passing it through several changes of pure wax inside the paratiin bath for
a few hours before embedding. If the material became brittle a shorter time in
the solution of clearing agent and wax was allowed. I have successfully cut
lizards’ ova § mm. in diameter when these precautions were taken.

When staining,’if the yolky sections washed off the slides this was usually
due to imperfect floating out of the wax ribbons on the slides, and imperfect
drying of the ribbons after floating out. The floating out was most successful
when done gradually and at a moderate temperature. Floating out on the top of
a paraffin bath is not a perfect method as the copper bath is usually much too
hot and the floating out dangerously rapid. I use a large piece of thick plate
glass arranged at a suitable distance above an electric light bulb, so that the glass
is just warm and the slides may be left on it for hours without damage, until they
are thoroughly dry. This method of dry heating eliminates the danger of imperfect
fixation due to a moist atmosphere.

It was found more important to take these precautions than to use a greater
quantity of egg albumen, since ribbons that are well floated out and thoroughly
dried should not leave the slides even when passed into 0:5% acid alcohol.

For an examination of anatomy the material was stained in bulk in carmalum.
For histological work, sections were stained in Delafield’s haematoxylin and
counterstained in eosin.

III. DESCRIPTION OF MATERIAL.

The material is described in four parts. The first part covers the period of
“pro-oestrus’’, the second the period of ovulation and fertilization, the third the
period of placental activity and the fourth the period after birth. The period of
placental activity is divided into three stages, Stage A presenting the early develop-
ment of the placentae and covering the first two weeks of the gestation period;
Stage B presenting the mature placentae, and covering the following eight weeks
of the gestation period; and Stage C presenting the placentae immediately prior
to the birth of the foetus and covering the last two weeks of the gestation period.

The description of the periods of ‘‘pro-oestrus” and of ovulation and fertiliza-
tion are based on examinations of lizards collected at sea level. However it is
more than probable that during these periods the condition of the reproductive
organs of females inhabiting the mountain regions is the same.

The descriptions of the period of placental activity and of the period after
birth are based upon an examination of both types of lizard, and as the placentae
of both are identical no distinction is made between them.

1. Period of “pro-oestrus’’.
As mentioned above, females, which presumably had not copulated, were
collected at Sydney on 10th September, 1926. The reasons for assuming that
copulation had not taken place are: when collected the ova were still within

BY H. CLAIRE WEEKES. 503

the ovaries and had not reached maturity, the largest ovum measuring 6 mm. in
diameter, the measurement at maturity being approximately 13 mm.; also, during
the beginning of September the weather is not warm enough to bring the lizards
out in any number, only three being caught on 10th September after searching
for two days; again, fertilization does not take place until the middle of October
and one would expect copulation to occur nearer the time of fertilization than
10th September, which is five weeks earlier, and although actual copulation was
not observed during the middle of October, it is extremely probable that it occurred:
then, since the lizards were observed living in pairs, a male with a female, the
females being easily distinguished by their greater girth due to the presence of the
enlarged ovaries.

The female reproductive organs consist of a right and a left ovary each equally
well developed, and two oviducts which open separately into the cloaca. Each
ovary is situated at the middle of the length of the corresponding oviduct. In each
of the two females collected on 10th September, the right ovary was more
anteriorly situated than the left, and the stomach was on the left side above the
left ovary and closely pressed against it. In one female there were four large
ova in each ovary, in the other there were four large ova in the right ovary
and three in the left. This species of lizard has only one breeding season each
year and the development of the ova is regulated accordingly, so that there is
little gradation in the size of the ova at this stage, but a marked contrast
between a number of small ova, all at approximately the same stage of development,
to a number of larger ova, also at approximately uniform stages in development.
At this stage these large ova have the same structure as that described for the
developing ova of Lacerta agilis (Hett, 1924), where the follicle cells are of two
varieties, some being enormously enlarged and the others comparatively small.

The oviducts are pleated, twisted and flattened against the dorsal body wall
of the lizard, the average length of an extended oviduct being 5 cm. The width
varies considerably, from 5 mm. to 2-5 mm. owing to an indication of division
into “incubatory chambers’’, which is due to the failure of the oviducts to regain
completely their natural shape after the preceding year’s gestation period.
Giacomini (1891) wrote that these chambers are evident in the oviducts of
C. tridactylus three months after birth. In ZL. quoyi it seems that they never
completely disappear.

The structure of the oviducts in these females is regarded as normal and only
variations from it, found in the oviducts during the period of placental activity,
are considered placental modifications. In section the wall of the oviduct at this
stage is more or less uniform in structure throughout its length with the exception
of the extreme anterior end which is modified for the reception of the ova. The
arrangement of the tissues of the oviductal wall seems uniform for lizards in
general, as it occurs in every species examined by the writer as well as in
C. tridactylus. The wall at the anterior end is thrown into folds, is comparatively
thin and the epithelium lining the lumen of the oviduct in this region is glandular
and ciliated, consisting of deep columnar cells which are much larger than those
lining the rest of the lumen. The wall of the remaining part of the oviduct
(Text-fig. 1) consists of the following layers: an external layer of peritoneum; a
thick muscular coat consisting of an outer coat of longitudinal muscle and an
inner coat of circular muscle; a thick mucous membrane in which glands are
embedded and which is lined by ciliated epithelium. The glands embedded in the
mucosa are simple or branched and open into the lumen by short narrow mouths.

M

504 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

EY

They are saccular, the cells being composed of vacuolated cytoplasm with small .
oval nuclei arranged round the periphery, and the cell walls being usually
indistinct. Many of the nuclei are undergoing mitosis, indicating the growth
of the glands. The epithelium lining the lumen of the oviduct is composed of
narrow ciliated cells, closely crowded together, with many of their nuclei also
dividing.

li

Ve — =) FF
<2 =a [-®
z

a ef

|
CIR. MUS. | LONG. MUS.

Text-fig. 1. Section of the wall of the oviduct of a non-pregnant female.
CAP., capillary ; CIL. UT: EPI., ciliated uterine epithelium; CIR. MUS., circular
muscle; GL, gland; LONG. MUS., longitudinal muscle. x 706.

The active cell division in the epithelium and glands indicates a “pro-oestrus”’
condition. The ordinary stimulus is certainly not copulation, since the ova in the
females under discussion are at a stage about five weeks prior to ovulation, and
it is extremely unlikely, as indicated above, that copulation has taken place. It is
more likely that the stimulus is an external periodical one in which temperature
is the main factor. That the breeding season of L. quoyi is influenced by climatic
conditions is evident from the fact that the lizards inhabiting the warmer regions
at sea level, Sydney, are at least two weeks earlier in their sexual season than
those inhabiting the colder regions at Mount Kosciusko, 300 miles south of
Sydney and with later seasonal changes.

2. Period of ovulation and fertilization.

On the 7th October, females containing ova at a stage prior to ovulation were
collected from the same locality as on the 10th September. The ova had reached
their maximum size and were packed with yolk. The membranes correspond with
those described by Hett (1924) for Lacerta agilis at this stage, the cells of the

BY H. CLAIRE WEEKES. 505

follicular epithelium being uniform in size and much smaller and flatter than
those surrounding the younger ovum described above. There is a thicker band
of thecal connective tissue in which numerous large blood vessels are present.

The oviducts have reached the final stage of preparation for the entrance of
the ova. Each is roughly divided into three regions, namely, the anterior end
described above, the more extensive middle region containing the majority of the
glands, and the extreme basal region leading to the cloaca containing no glands
and having its wall deeply convoluted. In section these regions differ somewhat
from the corresponding regions of the oviducts of females collected four weeks
earlier (10th September). At the anterior end the epithelial cells lining the
lumen of the oviduct are markedly glandular and their free surfaces are covered
by a thick secretion obviously derived from them and serving to facilitate the
entrance of the ova. In the middle region the glands are most numerous and
their cells are full of secretion and are distended until the central cavity is
obliterated, thus indicating a period of glandular activity. The nature of the
secretion from these glands is not apparent. At first, from the abundance of the
glands and their position in the middle region of the oviduct, it was naturally
supposed that they were for the secretion of albumen, but an examination showed
that there is no sign of albumen surrounding the ovum after its passage into the
oviduct or at any stage in its development. With the stretching of the “uterus”
upon the entrance of the ova and the consequent squeezing of the glands, their
secretion is forced out into the surrounding tissues of the uterus and some of it
passes between the muscle layers and in stained material is identical in appearance
with the substance of the shell membrane. The shell membrane is occasionally
found adhering to the mouth of one of the glands when it has been torn away
from the rest of the uterine wall, and a substance similar to it is present in the
mouth of the gland. The cells of the epithelium lining the lumen of the oviduct
in this region are also full of secretion, their cilia being matted together by the
exuded secretion which is thought to be for the general purpose of lubrication.

It is not definitely known to the writer whether the process of ovulation in
L. quoyi depends on some physiological stimulus such as the act of copulation, or
whether it is independent of such stimulus; also, whether the liberated ova pass
into the body cavity and thence into the oviduct with the aid of ciliary action, or
whether they pass directly into the oviduct as a result of the latter actively clasping
the ovum while still in the ovary.

As in other animals it is usual for the ova in each ovary to pass into the
oviduct on the corresponding side, but a female was collected which had received
an injury on the left side, damaging the left oviduct, so that, when ovulation took
place, the mature ova, five in number, passed into the right oviduct. The liberation
of ova from both ovaries was indicated by the presence of two burst follicles in the
left ovary and three in the right. The five ova in the right oviduct were so
tightly squeezed as to bend the oviduct completely out of position. The passage
of ova across the body cavity is not uncommon among mammals, having been
recorded for mammals ‘‘with a bicornuate uterus becoming pregnant in the uterine
horn on the side opposite to that on which the ovary had discharged’”’ (Marshall,
1910, p. 136).

When the ovum enters the oviduct and is fertilized, it becomes surrounded by
a thin shell membrane which is divided into three layers composed of matted
fibres. The uterus surrounds each egg as an expanded chamber called by Giacomini
the “incubatory chamber”. The wall of each incubatory chamber is uniform in

506 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

thickness measuring approximately 0-025 mm. and hence being much thinner than -
the wall of the uterus of a non-pregnant female. This decrease in thickness is
due to the stretching of the uterus on the entrance of the eggs. Owing to this
stretching, the coats of longitudinal and circular muscle are compressed into a
thin band of tissue and the glands, which are swollen and saccular in the uterus of
a non-pregnant female, are greatly compressed at this stage.

The epithelial cells lining the mucosa are much larger and not as crowded as
in the non-pregnant condition, and are of uniform size over the entire area of the
incubatory chamber.

There is a curious substance surrounding the ovaries after ovulation, which,
in prepared sections, has the appearance of a deep blue coagulum mixed with blood
clot containing corpuscles and numerous small round cells.

Immediately after ovulation the ruptured follicles are visible as large white
flat oval sacs, each with the cicatrix present as a median longitudinal groove.
However, a few days after ovulation they become smaller, spherical, yellow and
richly vascular, the alteration in the appearance of the follicles being due to the
presence of a corpus luteum in each, the growth of which is rapid, each follicle
being completely filled with luteal cells a few days after ovulation. It is not
probable that the corpora lutea have any influence on the retention of the ova of
reptiles comparable with their supposed function in mammals, since they occur
in ovaries of the oviparous lizard Lacerta agilis, which lays its eggs immediately
after their passage down the oviducts (Hett, 1924).

3. Period of placental activity.
Stage A. Early development of the placentae.

Twenty females were collected containing young embryos with placentae at
early stages in development, nine at Mount Kosciusko, eight at Sydney and three
at Kiama. Of the nine from Mount Kosciusko, seven contained three young, one
five and the remaining one six; of the eight from,Sydney, four contained six, two
seven and two eight; of the three from Kiama, one contained six, one seven and
one eight. In every case where a female contained an even number of young,
half were in each oviduct, and where a female contained an odd number of young
the right oviduct held one more than the left. This arrangement of embryos was
found to be the same in all lizards collected during the rest of the gestation period.

In the paper on the placentation in L. entrecasteauxi (Harrison and Weekes,
1925, p. 472) the authors wrote that “it seems remarkable that the number should
be odd in every one of nine examples, and we cannot find any explanation for this
condition”, also, “of the six females examined, four had more embryos in the
right oviduct than in the left’. Taking into consideration the conditions in both
L. entrecasteauxi and L. quoyi, it seems that there is a tendency for the female
to contain an odd number of embryos, and for the right oviduct to contain more
than the left. This may be due to the fact that the stomach of the non-pregnant
female is almost invariably on the left side and closely pressed against the left
ovary thus possibly interfering with the number of ova developing in this
ovary.

The developing embryo is dorsal in position with regard to the parent and lies
with its head directed mesially whether the embryo be in the right or left
oviduct. The embryos in the one female are not all at identical stages of develop-
ment, but the range of difference is negligible, L. quoyi resembling L. entrecasteausi
and not @. tridactylus in this respect. The uterus surrounds each egg as a thick

BY H. CLAIRE WEEKES. 507

white envelope which persists as the expanded incubatory chamber on the
extraction of the egg. These chambers are connected each to each by a short,
narrow, strap-like portion of the uterus which is deeply folded. Occasionally
the uterus is pigmented, the pigment being sometimes scattered over the surface
of each incubatory chamber, but usually restricted to the dorsal body wall of
the parent. The uterine wall shows as a smooth, semi-transparent membrane
whose uniformity is broken only by its thick opaque blood vessels, since there
are no villous foldings such as occur in C. tridactylus and L. entrecasteauzi.
However, when viewed through the binocular microscope the uterine wall is
seen to be covered by numerous branched glands (Text-fig. 2, A and B), which
are homologous with the saccular glands in the uterine wall of the non-pregnant
female (described above). They are present throughout the development of the
embryo and are outstanding and characteristic.

The vascularization of the wall of the incubatory chamber of L. quoyi is on the
same plan as that of L. entrecasteauzi. A single large artery and vein run
longitudinally along the dorsal wall of each uterus, the artery giving off branches
which pass transversely round the uterus to the base of the yolk-sac of the
contained blastocyst, where they break up into a rich network of capillaries,
and the vein receiving branches which also pass round the incubatory chamber
from the base of the yolk-sac, parallel to and roughly alternating with the arteries
(Text-fig. 2B). The villous folds in the wall of the uterus of L. entrecasteauzi
are fed by short branches from the main artery and vein and by branches from
the branch arteries and veins. In L. quoyi the allantoplacental region is similarly
vascularized.

In the early stages of development the uterus fits closely round the contained
embryos and keeps them in a fairly steady position, but with the preparation for
allantoplacentation a more perfect state of fixation results when the cells of the
chorion attach themselves to the epithelium of the uterus. However, before this
occurs the intervening shell membrane must disappear and in early stages of
development the chorionic cells appear to attack and absorb it. When the allanto-
placenta is mature there are often areas where a thin remnant of membrane
can be detected between the uterus and the modified chorion. The expansion of
the growing embryo aids the chorionic cells in their destruction of the shell
membrane by causing it to break and gradually fall away from the sides of the
embryo taking up a position at the base of the yolk-sac in the form of a flat
fibrous pad. This may be the “nodule” which occurs at the base of the yolk-sac
of the embryos of C. tridactylus, and which Giacomini (1891) terms the vitelline
membrane.

In L. quoyi the vitelline membrane is delicate and almost imperceptible and
could not possibly be confused with the shell membrane.

Of the two placentae the omphaloplacenta is the first formed, since its
requisites from the embryo, namely the chorionic ectoderm and a vascularized
yolk-sac, are formed early in development, whereas the allantoplacenta, depending
as it does on the presence of an allantois of sufficient extent to lie immediately
under the chorion, is comparatively late in its development.

The Omphaloplacenta. The embryos which show the earliest signs of
omphaloplacental modification are approximately one week old (Text-fig. 3, A),
their average length being 5 mm. Each has turned on to its left side, its head
being flexed and its body slightly curved. The amnion is completed and the
allantois present as a small swelling at the posterior end of the embryo. There

508 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

are three gill slits and approximately thirty somites. The chorionic ectoderm
completely surrounds the blastocyst and the extra-embryonic mesoderm extends
over a small area at the surface of the yolk-sac, which is entirely lined by
endoderm.

Although there are indications of foetal omphaloplacentation at this stage,
the uterine wall is as yet unmodified. In all earlier stages the development of the
extra-embryonic tissues is normal, the chorionic ectoderm cells being small and
in all respects resembling those in the chorion of oviparous lizards, but at this
stage the chorionic ectoderm at the base of the yolk-sac is slightly modified with
the beginning of omphaloplacentation and the further growth of the extra-

Text-fig. 2A. Incubatory chamber cut in half horizontally. Dorsal view
of ventral half showing the termination of the uterine arteries and veins in the
region of the base of the yolk-sac of the contained blastocyst, x 7:5; GL., gland;
UT. ART., uterine artery; UT. V., uterine vein.

embryonic mesoderm is abnormal. In the former case, a few of the cells of the
chorionic ectoderm at the lower pole multiply and become enlarged until a small
area of enlarged cells is formed (Text-fig. 3B and 4).

Occasionally there is more than one centre of modification, but each is small
and they gradually join together to form a single area. As the development of
the embryo proceeds and the omphaloplacenta approaches maturity, the modification

BY H. CLAIRE WEEKES. 509

of the chorionic ectoderm spreads over the entire under surface of the yolk-sac,
until a continuous sheet of moderately large columnar cells is formed.

In the embryos selected for description the area of modified ectoderm measures
approximately 0-8 mm. in diameter by 0-1 mm. in height, being thickest at the
centre (Text-fig. 4). It consists of a mass of deeply staining cytoplasm containing
irregularly arranged nuclei, there being no definite cell boundaries present, even at
the edge of the area where it merges into the unmodified chorionic ectoderm which
covers the rest of the yolk-sac. The yolk-sac endoderm immediately overlies and
mingles with the modified ectoderm and consists of a similarly staining cytoplasm
with scattered nuclei. The cytoplasm of the ectoderm is denser than that of the

Text-fig. 2B. Ventral view of dorsal half showing the main uterine artery
and vein, x 7:5. BR. GL., branched gland; M. UT. ART., main uterine artery;
M. UT. V., main uterine vein.

endoderm, which is vacuolated and has the appearance of having been mixed with
a fluid which coagulated during the process of fixation (Text-fig. 4), and hence
the line of junction between the two cytoplasmic regions is evident. The nuclei
embedded in the ectodermal cytoplasm are large and irregular in shape, with no
outstanding characteristics, and are easily distinguished from the endodermal
nuclei, which are larger when healthy but which are mostly degenerating, many

PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

VIT.VES.

VS.END,

BS

AR.MOD OMP==

Text-fig. 3. A, embryo approximately one week old, x 21:5; B, transverse
section of blastocyst containing an embryo about one week old, showing the
position of the area first modified for omphaloplacentation, x 11; AR. MOD. OMP.,
area modified for omphaloplacentation; CH., chorion; EX. COEL., extra-
embryonic coelome; VIT. VES., vitelline vessel; Y.S., yolk-sac; Y. S. END.,
yolk-sac endoderm.

BY H. CLAIRE WEEKES. 511

having been reduced to scattered groups of granules. The shell membrane under-
lies the chorionic ectoderm and between them is a gap which is partly filled with
a coagulum stained deeply by haematoxylin, and thought to be maternal secretion
passed through the shell membrane. It is possibly the presence of this secretion
in the protoplasm of the yolk-sac endoderm which gives the latter its peculiar
appearance. The uterine wall is pressed closely against the shell membrane and,
as stated, shows no indication of placental modification.

In embryos taken from a female collected during the second week of pregnancy,
the chorionic ectoderm cells are modified over a considerable area of the yolk-sac,
but the original centre of modification is evident at the middle of the base of the
yolk-sac where the cells are especially large and where the plug of endodermal
cytoplasm is still present (Text-fig. 5, A; Pl. xxxviii, fig. 1). Some of the
chorionic ectoderm cells are enlarged more than others, but are only about one-
third the height they eventually attain. They are narrow but definite, with their
free surfaces rounded and often bulging into peculiar shapes (Text-fig. 5, A,
CH. ECT.; Pl. xxxviii, fig. 1). The nuclei vary in size sometimes almost filling the
cell and they are arranged at the cell bases, staining deeply and standing out from
the rest of the cell cytoplasm. The cells are closely attached to an underlying
layer of yolk-sac endoderm, the cells of which are smaller than the ectoderm cells
and lie at the bases of the latter with the appearance of dovetailing. They are so
closely attached to the ectoderm cells that, when the layer of ectoderm is torn away

SH.MEMB CH.ECT. COAG UT.EPI- ¥S.END. DEG.NUC.

Text-fig. 4. Section of the omphaloplacental region, Stage A, showing the
area of modified tissue, x 511. CH. ECT., chorionic ectoderm; COAG., coagulum ;
DEG. NUC., degenerating nucleus; SH. MEMB., shell membrane; UT. EPI.,
uterine epithelium; Y. S. END., yolk-sac endoderm.

N

512 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

from the overlying yolk-sac in the preparation of material for sectioning, the
endoderm cells come away with it. The structure of the mass of endodermal
cytoplasm at the approximate middle of the base of the yolk-sac is unchanged.

DEG. NUG.

= a =e eee Et ke
—— Sa en SS
( EE CPUC SS GD o7 >»? ASH eee) Ce WEE. SMa TAs Nelbnes ~ CI yyy

Text-fig. 5. <A, section of the foetal portion of the omphaloplacenta of
embryos collected during the second week of the gestation period, x 562;
B, section of the maternal portion of the omphaloplacenta, x 562; DEG. NUC.,
degenerating nucleus; CH. ECT., chorionic ectoderm; END. CYT., endodermal
cytoplasm; UT. EPI., uterine epithelium.

BY H. CLAIRE WEEKES. 513

The structure of the uterine wall has changed slightly in this region. The
epithelial cells have increased in length, are definite and columnar, the protoplasm
at the free margins staining deeply, but they are very narrow and crowded
together and have not the same regular appearance as in more advanced stages
in the development of the omphaloplacenta.

The layer of endoderm underlying the area vasculosa on the dorsal surface
of the sac differs from the rest, consisting of large columnar cells, each with a
definite boundary and with its base rounded and buried in the yolk. The cells
are vacuolated and contain large nuclei and many small yolk granules (Text-
fig. 6, Y.S.END.). This layer of endoderm is homologous with that underlying
the area vasculosa in chick embryos (Lille, p. 129). The boundaries of the
endoderm cells filling the rest of the yolk-sac are difficult to determine, their
cytoplasm being full of yolk granules and large vacuoles. It will be recalled that
the yolk-sac endoderm is closely attached to the overlying chorionic ectoderm,
and this close attachment plays an important part in determining the structure of
the omphaloplacenta by bringing about the modification in the growth of the
extra-embryonic mesoderm as mentioned above. The growth is normal up to a
certain stage, the sinus terminalis being arrested for a short time at the junction
of the chorionic ectoderm and the yolk-sac endoderm. The area vasculosa under
these conditions measures on an average 7 mm. and consists of two main vitelline
arteries and veins. But the period of quiescence in the growth of the mesoderm
is short and the omphaloplacenta does not function for long in a non-vascularized
condition. The mesoderm cells at the circumference of the area vasculosa divide
fairly rapidly, but so close is the connection between the yolk-sac endoderm and
the chorionic ectoderm that the cells cannot force their way between them, and
consequently their progress is delayed and a thick margin of mesoderm cells is
formed surrounding the area vasculosa, which in transverse section has the
appearance of a rounded nodule of small dark cells (Text-fig. 6, A, NOD.). Since
there is no longer a passage for the extension of the mesoderm between the
ectoderm and endoderm it is forced to dip into the yolk-sac and continue its
growth round the yolk-sac, not over its surface as is usually the case, but
embedded in its substance (Text-fig. 6, B and C; Pl. xxxviii, fig. 2).

As the mesoderm pushes into the yolk-sac endoderm it splits into somato-
pleural and splanchnopleural layers as it does in the normal condition when
forming the extra-embryonic coelome, and thus separates an outer layer of
endoderm from the main bulk of the yolk-sac endoderm. This outer layer is fairly
regular in width at the sides of the yolk-sac, measuring approximately 0:075 mm.
in some embryos. However, at the base of the yolk-sac in the region of the
persistent area of modified ectoderm and endoderm, the mesoderm may grow up
into the yolk-sac for a considerable distance and so cut off an outer layer of
endoderm measuring as much as 0-210 mm. in width. At this stage (one week after
fertilization) the mesoderm has commenced to dip into the yolk-sac for the short
distance of 0-615 mm. and the area vasculosa still measures approximately 7 mm.,
there being no sign of haematopoiesis in the mesoderm within the yolk-sac
(Text-fig. 6, C). Since the mesoderm does not entirely surround the yolk-sac
until the embryo is at least four weeks old, the vascularization of the yolk-sac is
not completed until then and will be described at Stage B.

The Allantoplacenta.—Although the development of an allantois of sufficient
extent to lie immediately under the chorion is necessary for the formation of the
allantoplacenta, the maternal wall becomes modified in anticipation of placentation

514 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

when the embryo is a week old (Text-fig. 7, A), at the time when the allantois is
but a small swelling at the posterior end of the embryo and the omphaloplacenta
first begins to function. It is not until the embryo is approximately two weeks

SOM MES.
CH.ECT.

Text-fig. 6. A, section of the yolk-sac at the edge of the area vasculosa
showing the formation of the nodule of mesoderm; x 159. B, section of yolk-
sac of an older embryo showing the downgrowth of the mesoderm into the sac;
%* 159. OC, the same, under higher power; x 511. CH. ECT., chorionic ectoderm;
D. MES. Y. S., downgrowth of mesoderm into the yolk-sac; HX. COEL., extra-
embryonic coelome; NOD., nodule; SOM. MES., somatic mesoderm; Y.S.,
yolk-sac; Y.S. END., yolk-sac endoderm; Y. GR., yolk granule.

BY H. CLAIRE WEEKES.

old that the chorio-allantoic membrane is formed and the allantoplacenta
established.

At a stage a week after fertilization the amniotic folds have met above the
embryo to complete the chorionic membrane and it is in this region that the
uterine wall is first modified. The modification is brought about by the multiplica-

UT. GL. MAT. CAP.

Text-fig. 7. A, section showing the modification in the maternal wall before
the formation of the chorio-allantoic membrane; x 560. B, section through the
blastocyst at the end of the second week when the chorio-allantoic membrane is
first formed; x 8-5. C, the embryo at this stage; x 13. ALL. allantois;
CH., chorion; CON., constriction; MAT. CAP., maternal capillary; SH. MEMB.,
shell membrane; UT. GL., uterine gland; Y.S., yolk-sac.

516 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

tion and expansion of a few of the maternal capillaries present in the mucosa
between the saccular glands and the epithelium, and by their attempt to invade
the overlying epithelium (Text-fig. 7, A). The invasion of the epithelium by the
capillaries has not advanced far, the epithelium being intact over the surface of
the uterus with no breaks in its continuity such as occur at the stage of placental
maturity. It will be recalled that the epithelium lining the incubatory chambers
is a single layer of fairly large columnar cells. The pressure on these cells in
the region of the invading capillaries causes them to become flattened, so that they
now measure about 0:016 mm., one-third of their original height. The compression
of the cells is followed by their degeneration, which is most marked in later
stages, the compressed cells at this stage retaining their individuality although
their cytoplasm may be slightly vacuolated. Since the embryo is but a week
old the shell membrane between the uterine epithelium and the chorionic membrane
is practically unchanged, although the chorionic ectoderm cells have attacked it
and are embedded in its substance. The chorionic membrane is normal in
structure, the ectoderm cells being small, flat and unmodified. The growth of the
allantoic vesicle and the formation of the chorio-allantoic membrane occur during
the following week, so that, when the embryo is two weeks old (Text-fig. 7, B)
the main requisites for the formation of an allantoplacenta are first acquired,
there being a uterine wall in close proximity to a well vascularized chorio-
allantoic membrane. The blastocyst from the coastal type of female containing a
two weeks old embryo measures approximately 1:3 cm. x 0-9 cm., and the contained
embryo 0-6 cm., but its body is so curved, the head and tail almost meeting, that
it is impossible to give exact measurements, the length given being the crown-
rump measurement. The head is large, the eyes are prominent, and the limb buds
present, the stage in development roughly approximating that of a four-day-old
chick embryo (Text-fig. 7, C). The embryo lies on its left side and is sunk down
into the yolk-sac, the top of the sac often being level with that of the embryo
(Text-fig. 7, B). The area vasculosa extends a little more than half-way round
the yolk-sac and its boundary is marked by a constriction of the sac, which can
be seen in the living as well as in the fixed condition (Text-fig. 7, B, CON.).

The allantoic vesicle measures approximately 0-4 cm. in diameter. As the
embryo grows the vesicle expands radially until its wall meets the upper surface
of the yolk-sac which automatically stops further progress, the vesicle only
expanding as the yolk-sac decreases in size. As yet there is no definite, long
allantoic stalk, the allantoic vesicle opening directly into the hind gut and being
vascularized by two main arteries and veins.

The area of allantoplacentation is not limited, placental modifications of
maternal and foetal tissues occurring over the entire area embraced by the
allantois (Text-fig. 8, A). As the embryo develops, its yolk-sac is gradually
absorbed and the allantoic vesicle expands filling the former position of the yolk-
sac. The expansion of the allantoic vesicle and consequent radial extension of
the chorio-allantoic membrane is accompanied by a corresponding extension of
the placental region. It is peculiar that the uterine wall is always modified for a
short distance beyond the limit of its proximity to the allantois, and is thus
always prepared for the growth of the allantois and the extension of the
placental region.

At this stage the uterus is distinct from the chorionic membrane even when
there is no shell membrane present to divide them, the chorionic cells not having
yet begun to attach themselves to the uterus.

ed

BY H. CLAIRE WEEKES. 517

(a). Maternal Portion of the Placenta.—The wall of the uterus is thinner in
the placental than in the non-placental region due to the degeneration of the
glands and muscle layers and the partial degeneration of the epithelium. The
decrease in the number of glands in this region is particularly apparent when the
uterus is viewed through the binocular microscope (Text-fig. 2, A). The few
glands present are so compressed as to have no resemblance to their former
shape, the cell nuclei being arranged in a flat ring, and the cell cytoplasm being

Bo su.meme coer cap ee

Text-fig. 8. A, section of the dorsal portion of a blastocyst during the third
week of the gestation period, showing the extent of the allantoplacenta and the
general disposition of the membranes. B, section through the allantoplacenta,
stage A; x 1685. ALL. END., allantoic endoderm; ALL. VES., allantoic vesicle;
CH.-ALL., chorio-allantoic membrane; CH. ECT., chorionic ectoderm; FOET.
CAP., foetal capillary; GL., gland; MES. CO., mesodermal connection; SH.
MEMB., shell membrane; UT., uterus; VAC. CYT., vacuolated cytoplasm.

518 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

intensely vacuolated. The degeneration of the glands is accompanied by a
deposition of pigment granules.

The vascularization of the uterus is richer than in the first week of pregnancy.
The large vessels are situated in the mucosa overlying the glandular region and
the capillaries are dispersed among the glands particularly above the epithelium.
Each capillary has a distinct endothelial lining containing smail dark nuclei. The
capillaries are mostly small and flat and are not as numerous as at later stages.
They contain a few elongated corpuscles each with a deeply staining ellipsoidal,
almost rectangular nucleus. The capillaries have progressed in their invasion of
the epithelium and appear to push against the overlying epithelial cells, but they
have not as yet reached the surface of the uterus (Text-fig. 8, B). The degenera-
tion of the epithelium is more marked than in earlier stages. The cell walls and
many of the nuclei have disappeared and the cell cytoplasm is well vacuolated.
A few healthy cells are present and each has a small oval granular nucleus.

The persistence of the shell membrane varies for different embryos, some at
this stage having no membrane, some having a thin remnant, and others having
quite an appreciable amount. The membrane when present is pressed between the
chorion and the uterine wall, but there is a tendency for it to break away from
the uterine wall and remain attached to the chorion, this being due to the
absorption of the substance of the membrane by the underlying cells of the
chorionic ectoderm. That the shell membrane is being absorbed by the chorionic
cells is obvious, since the chorionic cells are found embedded in its substance
and in stained sections the cell cytoplasm has the same appearance as the
membrane; also the membrane is present in some places and not in others, and
is thinner in the placental than in the non-placental region.

(6). Foetal Portion of the Placenta—There is little modification of the
chorio-allantoic membrane, possibly due to the restricting presence of the shell
membrane, since the chorionic ectoderm cells must first destroy it before they can
come into contact with the uterus. The chorionic ectoderm consists of a single
layer of small, uniform, tapering cells with no visible dividing walls (Text-
fig. 8, B). Each cell contains an oval nucleus with usually two bright nucleoli,
the nuclei staining deeper than those of the uterine epithelium and hence being
easily distinguished from them throughout the life of the placenta.

The chorio-allantoic membrane is well vascularized by a rich network of
capillaries underlying the chorionic ectoderm. The capillaries measure 0:025 mm.
in diameter, being much larger than the maternal capillaries, and are full of young
corpuscles which are easily distinguished by their round shape and round nuclei
from the oblong maternal corpuscles. The connective tissue in which the vessels
and capillaries are carried is comparatively thick, consisting of a network of
young tapering cells, there being as yet no muscle fibres present among them.
There isa single bounding layer of allantoic endoderm, the cells of which resemble
those of the connective tissue. The restrictions to the efficient functioning of
the allantoplacenta are the presence of the shell membrane and the absence of any
modification of foetal tissue in the placental region. However it is evident that
a certain amount of maternal secretion is being absorbed by the foetal tissue,
since in this region there is none of the secretion from the compressed glands
in the mucosa such as is present in the non-placental area, described below. That
the chorionic ectoderm cells are ingesting is shown by their absorption of the
shell membrane and it is probable that at the same time they receive maternal
materials through it.

BY H. CLAIRE WEEKES. ; 519

The maternal portion of the placenta at this stage, then, is represented by a
general thinning of the uterine wall due to a partial degeneration of the muscle
layers and glands; by increased vascularization; by the invasion of the uterine
epithelium by the overlying capillaries and by the consequent compression,
flattening and degeneration of the epithelial cells. The foetal portion of the
placenta consists of a thin chorion attacking and absorbing the shell membrane
and overlying a well vascularized allantois.

INV. MAT. CAP.
UT. EPI.

CH. ECT.

EX.COEL.

Text-fig. 9. Section of the region of partial maternal placentation; x 1685.
CH. ECT., chorionic ectoderm; EX. COEL., extra-embryonic coelome; INV.
MAT. CAP., invading maternal capillary; UT. EPI., uterine epithelium.

(c). Region of Partial Maternal Placentation.—It will be recalled that the
uterine wall overlying the yolk-sac at the edge of the allantoplacental area is
influenced for some distance by allantoplacentation and is modified accordingly
(Text-fig. 9). Here the uterine wall is thicker than in the placental region and
the glands although still flattened are more prominent. The secretion which is
forced from these glands on the stretching of the uterus is present in abundance
among the tissues of the mucosa (Text-fig. 9). The uterine epithelium is thick
and the cells uniform, with indefinite boundaries, yet the capillaries have begun
to invade them, this invasion being the main placental modification. The shell
membrane is thick but is degenerating apparently through the activity of the

520 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

underlying chorionic ectoderm cells, which, however, are more easily separated ©
from it here than in the allantoplacental region. The chorionic ectoderm
immediately overlies the extra-embryonic mesoderm as far as the sinus terminalis,
when the mesoderm dips down into the yolk-sac as described, the chorionic
ectoderm then being attached to the endoderm cells of the yolk-sac. That the
uterine wall is secreting materials is quite possible but the presence of the
glandular secretion among the tissues of the mucosa in this region and its absence
in the placental region indicates the contrary.

Stage B. The Stage of Placental Maturity.

The stage of omphaloplacental and allantoplacental maturity covers the last
two months of the gestation period. The mature omphaloplacenta is not uniform
in structure, being more specialized in some embryos than in others and even
in its most specialized form is not as well developed as that in specimens of
L. entrecasteauxi, T. scincoides and EH. whitei examined by the writer. The
difference in the degree of specialization of the placenta makes it difficult to
estimate the exact time of its maturity, since the placentae in embryos at the
same stage in development may vary in structure, but the third week is thought
to be the most probable time of omphaloplacental maturity. The mature allanto-
placenta is more uniform in structure and the time of its maturity is conse-
quently more easily ascertained. The allantoplacenta is fully specialized to
function as a nutritive and respiratory organ when the embryo is between three
and four weeks old, and so functions with slight alterations in histological
structure over the ensuing two months prior to the birth of the foetus. The
structure of the placenta. of a fully formed foetus is influenced 1 approaching
birth and will be described separately as Stage C.

Nine females containing embryos with a mature omphaloplacenta and allanto-
placenta were collected from Barrington Tops at intervals during January and
December, 1925; twelve from the Blue Mountains during January, 1927, and
eight from Sydney during November and December, 1926. The embryos in the
females collected from Barrington Tops ranged in age from four to nine weeks;
those in females from the Blue Mountains from about nine to ten weeks; and
those in females from Sydney from three to ten weeks.

The difference in size between the blastocyst from the mountain type of
female and that from the coastal type is most noticeable, the blastocyst of the
mountain type measuring on an average one-half that of the coastal type. The
number of young present in each female corresponds with the figures quoted for
stage A, the females from the mountain region carrying from two to five young
and those from the coastal region from five to nine young. When there is only
a moderate number of blastocysts within the uterus and each embryo has room
to grow without interference from the others, the blastocyst is roughly oblong in
shape with its long axis parallel to that of the female, and each is isolated in its
incubatory chamber; but when they are crowded and pressed together their
depth is as great as, if not greater than, their length. They are sometimes
pressed so closely together that the uterus is no longer constricted between them
to form definite incubatory chambers but remains distended so that the embryonic
membranes over a small area at both ends of one blastocyst are in close proximity
with those of the two adjacent blastocysts.

As in L. entrecasteauxi (Harrison and Weekes, 1925, p. 473) certain structures
are more or less visible through the thin and much distended uterine wall, and

BY H. CLAIRE WEEKES. 521

“as the parent lies upon her back, with the ventral body wall opened up, and the
uteri exposed, the region of the yolk-sac for each embryo shows as a creamy
area, ventral and lateral, i.e., anti-mesometrial in position’. The placentae of
embryos obtained from females during the fourth week of the gestation period
are selected for description and such variations as are found in the placentae of
embryos at later stages will be described separately. When the four weeks old
blastocyst is examined ventrally, although the yolk-sac has been absorbed from
the immediate region of the embryo and reduced to half its original size, it
obscures from view most of the underlying embryo, which is only completely
visible when the blastocyst is turned round. The embryo can then be seen lying
on its left side on the yolk-sac with its body curved so that head and tail meet,
and with the tail coiled round the limbs. The vascularization of the uterine wall,
allantoic vesicle and yolk-sac can be clearly seen. Beyond a general enlargement of
the vessels in the uterine wall the maternal circulation does not differ from that
described at stage A, the vessels encircling each incubatory chamber as far as
the base of the yolk-sac.

LUMB, VES IN. ALL.MEMB. BST. Y.S.ST. © ALL. ST.

Text-fig. 10. Four weeks old embryo lying upon the yolk-sac, showing the
general disposition of the foetal membranes; x 7:5. ALL. ST., allantoic stalk;
B. ST., body stalk; C. BR., cellular bridge; IN. ALL. MEMB., inner allantoic
membrane; L. UMB. VES., left umbilical vessels; Y. S. ST., yolk-sac stalk.

The embryo at this stage (Text-fig. 10) is closely wrapped in the amnion and
is a uniform white colour, there being no scale marking nor any indication of scales
in sectioned material. The head is large, the eyes prominent, the nasal apertures
formed, and the mouth distinct with upper and lower jaws, tongue and rudimentary
teeth forming. The joints and digits of the fore and hind limbs are fairly well
developed. The body stalk leaves the posterior end of the embryo in the region
of the hind limbs and soon separates into yolk-sac and allantoic stalks. The yolk-
sac stalk passes immediately downwards carrying the vessels connecting the
vascularization of the yolk-sac with that of the embryo. The allantoic stalk
passes up and round the embryo expanding into the allantoic vesicle which

522 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

measures approximately 5 mm. if the embryo is from a female of the mountain
type and 13 mm. if from one of the coastal type. As in earlier stages the
allantois does not surround the yolk-sac and only expands in a downward direction
as the yolk-sac decreases in size.

In the body stalk the allantoic stalk has the appearance of being folded round
the yolk-saec stalk, owing to the presence of allantoic vessels on either side of the
embryo, while the yolk-sac receives vessels from one side only. The yolk-sac stalk
is narrow in comparison with the allantoic stalk and its cavity has almost
disappeared, being lined with fairly large columnar cells (Text-fig. 11, A). The
cavity of the allantoic stalk is much larger and is also lined with large regular
cells. The yolk-sac stalk enters the gut immediately anterior to the allantoic stalk.
The various bloodvessels are slung in thick folds of mesenchyme arranged round
the two stalks, the mesenchyme of both stalks intermingling. There are two
umbilical arteries and two veins arranged round the cavity of the allantoic stalk,
and one vitelline artery and one vein round the cavity of the yolk-sac stalk. In
Text-fig. 11, A, the relationship between the allantoic and oe -sac stalks within
the body stalk is shown.

Within the body of the embryo the vitelline artery is given off by the main
dorsal aorta, and the vitelline vein enters the liver. Hach umbilical artery is a
separate branch from the dorsal aorta in the region of the sciatic artery. As the
umbilical veins enter the embryo they unite and pass to the liver where they join
the intra-hepatic vessels and the ductus venosus and so pass to the heart. The
vitelline artery and vein branch over the flat upper surface of the yolk-sac, at the
edge of which they dip into its substance (as described at Stage A), and so
completely encircle the sac. The upper surface of the yolk-sac is covered with a
layer of large compact endoderm cells (described at Stage A) into which the
vessels sink until they lie in deep grooves. The cells lining the grooves prevent
the vessels from penetrating into the interior of the yolk-sac and it is not until
the vessels have passed beyond the region of these cells and have reached the
sides of the yolk-sac that penetration of the interior is possible. It is curious that
the interior of the yolk-sac is vascularized by the vessels from the sides and base
of the sac and never by branches of those on its upper surface. However, the
large endoderm cells are thought to be probable yolk absorbers, since they retain
their characteristic appearance throughout the life of the yolk-sac and are always
packed with minute yolk granules.

The area vasculosa extends right round the yolk-sac and so the extra-
embryonic coelome is completed. It will be seen at once that the position of the
coelome is abnormal, since it actually lies within the yolk-sac, and the somato-
pleure is formed from mesoderm, endoderm and ectoderm instead of from the
normal layers of mesoderm and ectoderm. At the edge of the yolk-sac the somato-
pleural mesoderm lining the chorionic ectoderm remains attached over a short
distance to that of the splanchnopleure of the yolk-sac, and the mesoderm of the
allantois joins this mesenchyme connection, so that the extra-embryonic coelome is
divided into two parts, one lying in the embryonic region between the chorionic
membrane and the amnion and being almost completely filled by the allantoic
vesicle, and the other lying within the yolk-sac. The attachment of the allantois
to the chorionic and yolk-sac mesoderm in this region was detected when the
embryonic membranes were dissected away from the embryo under the binocular
microscope and was puzzling until sections were made which showed its nature.
The blood is collected from the chorio-allantoic membrane by the two large

BY H. CLAIRE WEEKES. 523

umbilical veins one of which passes around the inner allantoic wall and thence
to the allantoic stalk; the other, invested by allantoic mesoderm and endoderm,
reaches the allantoic stalk by passing directly across the cavity of the allantoic

ALL.ST.

UMB. Aw
: VIT. ART.

VIT.V.

Text-fig. 11. A, section of the body stalk showing the general relationship
between the allantoic and yolk-sac stalks; x 86. B, section of the allantoic
vesicle in the region of the vascular bridge; x 16:5. ALL. ST., allantoic stalk;
ALL. VES., allantoic vesicle; C. BR., cellular bridge; CH. ALIL., chorio-allantoic
membrane; IN. ALL. MEMB,., inner allantoic membrane; UMB. ART., umbilical
artery; UMB. V., umbilical vein; UT., uterus; VIT. ART., vitelline artery;
VIT. V., vitelline vein; Y. S. ST., yolk-sac stalk.

524 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

vesicle (Text-fig. 11, B). Each of the veins is accompanied throughout its.
ramification in the chorio-allantoic membrane by one of the umbilical arteries.
This method of transmission of blood vessels across the cavity of the vesicle is
interesting but is evidently of common occurrence since it is found in embryos of
some mammals and reptiles examined. Hubrecht (1889, pp. 307-8) records it for
the hedgehog, Flynn (1923, p. 77, and fig. 1, p. 74) for Tiliqua scincoides, and
Harrison and Weekes (1925, p. 476) for L. entrecasteauxi. For the hedgehog
Hubrecht (1889, pp. 307-8) described “cellular bridges” which crossed the allantoic
cavity to carry blood vessels from the inner to the outer allantoic membrane.
Flynn wrote, “But the outer wall of the vesicle is also supplied by vessels which
leave the allantoic stalk near the body of the embryo and pass right across

Text-fig. 12. Diagrammatic representation of the formation of the vascular
bridges across the allantoic cavity. ALL. CAV., allantoic cavity; ALL. F.,
allantoic fold; ALL. VES., allantoic vessels; JUN. ALL. F., junction of allantoic
folds; PL., pleat.

BY H. CLAIRE WEEKES. 525

the vesicle to ramify through the mesenchymal layer of the placental face of
the allantois’. Harrison and Weekes (p. 476) described a somewhat different
method of transmission for L. entrecasteauxi where “a pleated fold arises from
the inner allantoic wall (Plates xlvii, fig. 3; xlviii, fig. 2; xlix, fig. 5, Pl.), within
the inner free edge of which the vessels are carried. Since this pleat is covered
externally with endoderm continuous with that lining the allantois, there would
appear to be no doubt that it has arisen originally as a fold of the allantoic wall”.
The authors also noted that “In our fourth series a somewhat different condition
occurs. In place of the flat fold a blunt finger-like process is pushed out from
the inner wall and passes across the lumen to bring about the same ultimate
result”.

An examination of the various available early stages in the development of
L. quoyi reveals the suprisingly simple method of the formation of the bridges
across the allantoic cavity and indicates a similarity between the methods of
transmission in L. quoyi, L. entrecasteauxi and T. scincoides and possibly in the
hedgehog.

The position of the. blood vessels in the allantoic vesicle in L. quoyi is due
to the comparatively rapidly growing wall of the vesicle enfolding the more
slowly growing blood vessels present in its mesenchyme (Text-fig. 12B, ALL. F.).
The arms of the folds from either side meet (Text-fig. 12, C), and thus separate
the vessels from the rest of the vesicle wall except for a pleat-like connection
which may remain helping to hold the suspended vessels in position (Text-fig.
12, D, PL.). Such a pleat may evidently persist in L. entrecasteauxi as quoted
above, but in LZ. quoyi, in.one of the four specimens of L. entrecasteauxvi examined,
and apparently in 7. scincoides, it becomes disconnected from the main vessels
which then have the appearance of being carried across the cavity in “blunt
finger-like processes” pushed out from the walls of the vesicle. The enfolding
of the vessels by the wall of the vesicle accounts for the presence ef the coat of
allantoic endoderm surrounding them, and, I think, for the substance of the
“cellular bridges’ in TJ. scincoides mentioned by Flynn, although he has not
indicated the nature of their tissue either in the text or the accompanying figure.

Since the embryo lies on its left side, the underneath wall of the allantoic
vesicle is pressed against the posterior end of the embryo and the yolk-sac and is
so prevented from expanding in a downward direction; consequently the lower
wall of the vesicle, which is vascularized by the left umbilical vein and artery,
does not grow to the same extent, nor as rapidly, as the upper wall, and this, I
think, accounts for the absence of “cellular bridges” carrying the left umbilical
vein and artery across the cavity of the allantoic vesicle. The inner allantoic
wall is so tenuous and so closely wrapped round the tail and hind limbs of the
embryo, that in serial sections one may easily be misled into believing that the
allantoic stalk passes downward and outward over the surface of the embryo.
However, a glance at Text-fig. 10 will show that sections cut along the plane x-x',
in the region of the left umbilical vein and artery, would have this appearance.
In Text-fig. 10 the embryo is drawn lying upon the yolk-sac with the body stalk
.(B. ST.), leaving the posterior end of the body and dividing into the yolk-sac
stalk (Y.S. ST.), and the allantoic stalk (ALL. ST.). A portion of the wall
of the allantoic vesicle has been removed so that the transmission of the blood
vessels across the allantoic cavity in the “cellular bridge’? can be seen (C. BR.)
as well as the folding of the inner allantoic membrane round the tail of the
embryo (IN. ALL. MEMB.), and the consequent passage of the left umbilical

525 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

vein and artery (L. UMB. VES.), to the chorio-allantoic membrane in a groove of.
the inner allantoic wall.

The Omphaloplacenta.—The lack of uniformity in the degree of specialization
of the omphaloplacenta in different embryos is, as stated, most probably due to
the varying amount of shell membrane present underlying the yolk-sac. It will be
recalled that the shell membrane falls away from the sides of the blastocyst and
collects at the base of the yolk-sac between it and the uterine wall. When the wall
of an incubatory chamber is cut in half round its circumference, and the lower
half is gently lifted from its position underlying the yolk-sac, the remnant of the
shell membrane in the form of a round flat pad of crumpled fibrous matter falls
away from the base of the yolk-sac, since it is no longer supported by the uterus.
The pad gradually thickens as the embryo grows and more membrane falls to
the base of the yolk-sac, so that, while it is comparatively thin in the majority
of embryos at four weeks, in embryos collected during the second and last month
of the gestation period, it is of considerable thickness, and impresses its shape
upon the overlying yolk-sac, forming a barrier between it and the uterine wall
and so interfering with the modification of the foetal tissue of the omphalo-
placenta. When the shell membrane is thin the foetal tissue reaches its highest
degree of specialization; when it is thick in some places and thin or absent in
others the modification of the foetal tissue is correspondingly patchy; and when
it is abundant, as in embryos collected during the latter half of the gestation
period, there is only slight modification of the foetal tissue. In L. entrecasteauxi
(Harrison and Weekes, 1925, p. 474) there was no trace of shell membrane in any
of the embryos examined, and consequently the placenta is more or less uniform
in its development and is more highly specialized than that of L. quoyi, even at
the late stage described when retrogression of its structure is suspected (p. 476).

The embryos collected during the fourth week of the gestation period have
the most fully specialized omphaloplacenta (Text-fig. 13). Here, although the shell
membrane completely covers the surface of the yolk-sac, it is thin and the pad
at the base of the sac comparatively small.

(a). Maternal Portion of the Placenta.—The maternal portion of the placenta
is much less influenced by the presence of shell membrane than the foetal portion,
and is comparatively uniform in structure throughout the period of omphalo-
placental maturity.

There is a tendency towards a thickening of uterine tissue at the base of the
yolk-sac which is fairly consistent in all embryos and especially marked in those
at four weeks. The increase in thickness is brought about by a thickening of the
muscular tissue and the mucosa, and by the enlargement of the epithelial cells.
There is a tendency toward slight folding of the uterus, the folds being irregular
and the crypts shallow. The glands in the allantoplacental region described at
Stage A are conspicuous throughout the mucosa over the placental area, but
although fairly numerous do not appear to function, since the cytoplasm of the
gland cells is vacuolated and the cells are arranged round a large central cavity.
The capillaries are numerous and mark the termination of the uterine vessels
which pass round the incubatory chamber (Text-fig. 2, A). They are flat, but
not as flat as in earlier stages (Text-fig. 5, B), and often bulgé beneath the
epithelium as in L. entrecasteauri. The thickening of the uterus restores to it
something of its appearance in the non-pregnant condition, but signs of placenta-
tion are present in the increased vascularization and the modified epithelium.
The epithelium is formed of fairly large columnar cells with oval nuclei and with

BY H. CLAIRE WEEKES. 527

the cell cytoplasm concentrated at their rounded free margins, this concentration
being probably due to the secretory activity of the cells, since a coagulum is
found adhering to many of them.

In some embryos at various stages in development the connections between
the incubatory chambers are so deeply convoluted that when met with in sectioned
material the uterine wall in these regions appears to be thrown into deep regular
folds, which may be at first thought to be a placental modification.

(b). The Foetal Portion of the Placenta—The modification of the foetal
tissue extends over the entire under surface of the yolk-sac in proximity with the
uterine wall by the radial extension of the original area of modified tissue at the
approximate middle of the base of the yolk-sac. At this stage the chorionic
ectoderm is thickest at the base of the sac and is composed of two or three
layers of cells. The lower layers of the ectoderm are flat and squamose, and the
difference between the outermost layer of cells and the underneath layers is

EX.COEL. SP.MES. Nes:

CH.ECT.

UT.EP] § SH.MEMB. Ge

Text-fig. 13. Section of the mature omphaloplacenta; x 103. CH. ECT.,
chorionic ectoderm; EX. COEL., extra-embryonic coelome; GhL., gland;
SH. MEMB., shell membrane; SP. MWS., splanchnic mesoderm; UT. EPI.,
uterine epithelium; Y.S., yolk-sac.

marked, the cells of the outer layer being tall and narrow with many of the
individual cell boundaries distinct. Although the majority of the cells are
columnar others may be crowded together until they have no definite shape. At
the sides of the yolk-sac the ectoderm is thinner and the cells of the outer layer
are smaller. There is a comparatively thin layer of shell membrane adhering to
the chorionic ectoderm over the placental face but it is being absorbed by the cells
and has no apparent influence on their development.

The yolk-sac endoderm underlying the chorionic ectoderm contains yolk-
granules and is composed of large, vacuolated cells with large deeply staining
nuclei. It is not so intimately attached to the chorionic ectoderm as in earlier

(0)

528 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

stages and the degree of attachment varies with the embryo. The boundaries of
the cells are not definite, and it is impossible to estimate the number of cells in —
the depth of the layer. The endoderm cells are bounded by a layer of non-
vascularized extra-embryonic mesoderm.

The chorionic ectoderm, together with the underlying yolk-sac endoderm and
extra-embryonic mesoderm, is separated from the main bulk of the yolk-sac, with
the exception of a small area at the approximate middle of the base of the sac
where the mesoderm remains attached to that lining the sac.

The maternal secretion absorbed by the chorionic ectoderm must be passed
through the narrow layer of yolk-sac endoderm and somatopleura!l mesoderm
and across the cavity of the extra-embryonic coelome before it reaches the blood
vessels in the yolk-sac. Hence the fine coagulum within the extra-embryonic
coelome, which contains corpuscles and cell debris, is thought to be maternal
secretion.

The omphaloplacenta, then, present in embryos four weeks old is fairly well
developed and is formed by a richly vascular uterine wall with epithelium
modified for secretion overlying a many layered sheet of modified chorionic
ectoderm cells, attached to a thick layer of large vacuolated endoderm cells
followed by a layer of non-vascular mesoderm; the sheet of combined foetal tissue
being separated by a narrow extra-embryonic coelome from the vascular system
of the yolk-sac. The extra-embryonic coelome is completely cut off from the rest
of the coelome in the embryonic region by a mesenchymal connection between
the allantois, chorion and yolk-sac.

In embryos obtained from females collected during the sixth and subsequent
weeks of the gestation period, the omphaloplacenta is not so well specialized.
The maternal placenta in’ these corresponds with that described above and needs
no further comment. During the sixth and seventh weeks, when there is a
patchy accumulation of shell membrane at the base of the yolk-sac, some areas of
foetal tissue are more specialized than others, the foetal placenta in the embryo
selected for description being roughly divisible into two regions of varied specializa-
tion. The first region covers about one-half of the yolk-sac extending from the
edge of the sac to its base and consists of practically unmodified foetal tissue
underlain by a convoluted layer of shell membrane. The second region covers
the rest of the yolk-sac, and as there is very little shell membrane here, the
foetal tissue is well modified, the chorionic ectoderm cells attaining a height of
0-087 mm. (Text-fig. 14, A).

The chorionic ectoderm in the first region consists of two or three layers of
cells which are small and flat with no dividing walls visible. The cell cytoplasm
stains a deep pink with eosin and the nuclei stand out as oval darkly stained
bodies. The free margins of the cells are irregular, since the cells are embedded
in the shell membrane which is surrounded and impregnated by a thick coagulum
thought to be maternal secretion.

The cells of the chorionic ectoderm in the second region are arranged in
many layers, those of the outermost layer ranging from short, wide, cubical cells
to tall, narrow, irregularly shaped cells with a tendency to columnar structure
(Text-fig. 14, A). These enlarged cells are phagocytic and have ingested material,
often uterine epithelial cells and maternal corpuscles, in their cytoplasm. Between
the uterine wall and the chorionic ectoderm there is the same pale staining
coagulum which occurs in the first region and which adheres to the edge of the
chorionic cells giving them a ragged appearance. This coagulum contains the

BY H. CLAIRE WEEKES.

Text-fig. 14. A, section of the foetal tissue of the omphaloplacenta of a six
weeks old embryo showing the irregularly shaped chorionic ectoderm cells;
x 1050. 3B, ‘finger-like’ downgrowth of chorionic ectoderm in the omphalo-
placental region of an eight weeks old embryo; x 1050. CH. ECT., chorionic
ectoderm; F. GR. CH. ECT., finger-like growth of chorionic ectoderm;
SH. MEMB., shell membrane; SOM. MES., somatic mesoderm; Y.S. END.,
yolk-sac endoderm.

529

530 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

scattered cells and corpuscles which the chorionic ectoderm cells ingest. The
underlying layer of yolk-sac endoderm is thinner than in the placenta of the four
weeks old embryo, the narrow cells tapering and occasionally bulging round
relatively enormous yolk granules. The extra-embryonic coelome is wide and in
some of the embryos examined is filled with a coagulum which is stained a deep
blue by haematoxylin and which in patches has a peculiar scale-like appearance.
It is full of corpuscles and has a fine granular matrix at its edges, which has
the appearance of congealed blood. It is not known whether the corpuscles are
maternal or foetal but both varieties are possibly present. Its occurrence between
the uterine wall and the chorionic ectoderm, within the yolk-sac and within the
allantoic vesicle as well as within the extra-embryonic coelome makes the problem
of its origin a difficult one. A similar coagulum was observed in the yolk-sac of
L. entrecasteauzi (Harrison and Weekes, 1925, Pl. xlviii, fig. 5) and in FH. whitei.
It was at first thought to be albumen, but this is not likely as it was not present
in young embryos examined nor in the majority of older ones. Some of it is
probably maternal secretion but it is not likely that this is the sole source since
there is such a large quantity of coagulum present. Owing to its blood-like
appearance it was thought that its presence may be due to the breaking of
vessels damaged when the parent lizard was caught (Pl. xxxix, fig. 3).

In embryos present in females collected during the last month of the
gestation period, the pad of shell membrane practically covers the under-
surface of the yolk-sac, and there is consequently only slight specialization of the
foetal tissues. For a considerable distance on either side of the yolk-sac the
chorionic ectoderm cells are normal, each being flattened and having a small dark
nucleus. In the central region where the shell membrane is thrown into deep
folds the chorionic ectoderm is formed of many layers of small cells. These
cells proliferate and grow down between the folds of the shell membrane in the
form of long “finger-like” growths which may measure as much as 0:25 mm. in
length and which are usually about three or four cells thick (Text-fig. 14, B).
The hollows between the folds of the shell membrane on the side nearest the uterus
are full of a thick coagulum which contains corpuscles and which stains pink in
some places and a deep blue in others (eosin and haematoxylin). It is most
probably maternal secretion and is thought to supply the stimulus for the down-
growth of the chorionic ectoderm into the folds of the shell membrane. The
cytoplasm of the cells of the proliferated areas has the same dense pink and blue
appearance as the coagulum which the cells are obviously absorbing. Over a
small area at the edge of the yolk-sac where the shell membrane is not folded
but is flat and comparatively thin, the chorionic ectoderm cells are much larger
and resemble the cubical chorionic ectoderm cells in the omphaloplacental region
in the six weeks old embryo.

The underlying layer of yolk-sac endoderm has the same general structure
as in the six weeks old embryo, being composed of either long tapering cells or
large vacuolated cells. Often for a considerable distance at one side of the yolk-
sac the chorionic ectoderm and strip of yolk-sac endoderm are attached to the
yolk-sac.

The omphaloplacenta, then, functions best during the first six weeks of the
gestation period when the foetal tissues are most highly specialized and the
shell membrane covers the yolk-sac as a smooth layer. During the latter half
of the gestation period its activity is diminished by the presence of the thick
pad of shell membrane underlying the yolk-sac, which interferes with the passage

BY H. CLAIRE WEEKES. 531

of maternal secretion from the uterus to the embryonic tissues, some of the
secretion being retained among the folds of the membrane.

The Allantoplacenta.—By the end of the fourth week of the gestation period
there is usually no shell membrane left and the chorio-allantoic membrane is
pressed closely against the smooth face of the uterus and the fixation of the tissue
is brought about by the attachment of scattered enlarged cells of the chorionic
ectoderm to the uterine epithelium. In the handling and sectioning of the
material, part of the foetal tissue may separate from the maternal tissue but
there are always large areas of foetal tissue left attached to the uterus. The
attachment though efficient is not as close as that in Perameles and there are
no villous folds in the uterine wall to aid in fixation.

(a). The Maternal Portion of the Placenta.—The uterine wall is about three
times thicker than in the allantoplacenta of a two weeks old embryo and the
placenta is uniformly developed over its entirety, there being no special thickening
of the uterine wall at the centre of the placental region such as occurs in the
omphaloplacenta. With the thickening of the uterus the muscle layers, the
mucosa and the few glands present have resumed something of their normal
structure, but the glands show no signs of activity. The numerous capillaries
have reached the surface of the epithelium and are now only separated from the
underlying foetal tissue by their own endothelial walls and perhaps a thin layer
of maternal cytoplasm (Text-fig. 15). They are crowded at the surface of the
epithelium with their walls often touching, but with usually the width of their
diameter apart. The capillary walls are thin but definite, with occasional small
sickle shaped nuclei present in their substance. The capillaries are circular,
packed with corpuscles, and are overlain by a series of larger vessels in the
mucosa which are often found pressing against the underlying capillaries.

VAC. UT. EPI. EPI. NUC.

ENL. CH. ECT. C. ALL. CAP.

Text-fig. 15. Section of the allantoplacental region of a four weeks old
embryo; x 1270. ALL. CAP., allantoic capillary; ENL. CH. ECT. C., enlarged
chorionic ectoderm cell; EPI. NUC., epithelial nuclei; GL., gland; VAC.
UT. EPI., vacuolated uterine epithelium.

532 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

There are few definite epithelial cells present, since most of them have
degenerated, the cell cytoplasm being full of small vacuoles, the cell walls having
disappeared, and a few of the degenerating nuclei being represented by groups
of small dark granules. The healthy nuclei are round, stain but slightly, have
one or two nucleoli, and are quite characteristic and always distinguishable from
the nuclei of the chorionic ectoderm. These nuclei have been pushed aside by
the invading capillaries and are now grouped in the cytoplasm between them.
The cytoplasm has not the appearance of that of typical secreting cells such
as are seen lining the villous ridges of L. entrecasteauxi, and hence is not
regarded as an active secreting agent.

The absence of shell membrane allows the chorionic ectoderm and the
uterine wall to come into immediate contact, but although they are attached,
the connection is such that the line of division between maternal and foetal
tissue is often visible, there being no mingling of the chorionic ectoderm with
the maternal tissues such as occurs in the placentae of some mammals. The
chorionic cells which attach themselves to the uterine epithelium are comparatively
widely separated and have no definite shape, consisting of a large deeply staining
nucleus surrounded by cytoplasm which insinuates processes into the maternal
cytoplasm, so that when the attached maternal and foetal tissues are separated
both surfaces are jagged and torn. For the rest the chorionic ectoderm consists of
long tapering cells with fairly large nuclei, the tapering ends of the cells
extending over the bulging allantoic capillaries or degenerating to allow the
capillaries to come into immediate contact with the maternal capillaries (Text-
fig. 15). It is difficult to determine whether there is more than one layer of
ectoderm cells present. In some places there is a narrow gap between the
maternal and foetal tissues which is filled with a coagulum staining lightly with

haematoxylin, and which is possibly maternal secretion, although it may be the
remains of shell membrane.

The allantoic blood vessels vary in size from small round capillaries to much
larger vessels. The capillaries have distinct nucleated walls and are filled with
corpuscles which closely resemble the maternal corpuscles at this stage. A
narrow band of connective tissue underlies the capillaries and contains less
muscle fibre than that in L. entrecasteauxi. The single layer of endoderm
bounding the chorio-allantoic membrane consists of fairly regular, moderately
enlarged tapering cells. At the edge of the placental area, where the outer wall
of the allantois bends at the surface of the yolk-sac and continues as the inner
wall, the endoderm cells may proliferate until they hang in bunches in the
allantoic cavity (Text-fig. 16, A). Here, cell boundaries are difficult to determine,
some of the cells appearing to be multinucleate. Each cell has a narrow base
and a swollen apex which often contains one or more large vacuoles. The
modification of the cells may extend over the inner wall of the allantois, where
the cells enlarge and separate from each other at their margins (Text-fig. 16, B).
They are often intensely vacuolated with a deeply staining nucleus at the
surface of the cell. In the allantoplacental region of some rodents the endoderm
cells have the same peculiar structure, but this modification is not a placental
adaptation as is shown by a similar modification of some of the endoderm cells
lining the allantoic vesicle of the oviparous Agamid lizard Amphibolurus barbatus.
The peculiar structure of the cells is thought to be possibly caused by their
absorption of the excretory fluid which fills the allantoic vesicle.

BY H. CLAIRE WEEKES. 533

The inner allantoic membrane may unite with the amnion and the junction
may extend completely over the surface of the latter, so that the blood vessels of
the inner allantoic membrane come to lie between a layer of allantoic endoderm
and amniotic ectoderm (Text-fig. 16, B). The ectoderm cells of the amnion
are also modified, their structure resembling that of the endoderm cells just
described, and this similar modification is possibly due to their absorption of
excretory fluid from the endoderm cells.

AREA NO1.

C

CH-ALL.MEMB.

AREA NO 2.

Text-fig. 16. A, endoderm cells lining the allantoic vesicle at the surface of
the yolk-sac; x 405. B, section showing the structure of the fused inner
allantoic membrane and amnion; x 405. CC, section of the allantoplacental
region and allantoic vesicle showing the junction of the inner allantoic membrane
and amnion with the chorio-allantoic membrane; x 17:5. ALL. CAV., allantoic
cavity; ALL. END., allantoic endoderm; AMN. ECT., amniotic ectoderm;
CH. ALL. MEMB., chorio-allantoic membrane; IN. ALL. MEMB., inner allantoic
membrane; UT. WALL., uterine wall.

In the majority of placentae examined an interesting condition exists, where
all the extra-embryonic membranes—the chorion, the outer allantoic and inner
allantoic membranes, and the amnion—join together in some places to form
the foetal portion of the allantoplacenta (Text-fig. 16, C; Pl. xl, fig. 7). In some

534 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

of the placentae examined the area so formed is comparatively large, measuring ©
as much as 5 mm. in one of the embryos obtained from a mountain type of female.
The presence of the additional membranes modifies the structure of the chorio-
allantoic membrane in their vicinity, as described below.

CH. ALL.MEMB.
AMN.ECT.

Text-fig. 17. Section of the first area of combined foetal membranes in the
allantoplacental region, showing the compressed chorio-allantoic membrane and -
well vascularized inner allantoic wall; x 519. AMN. ECT., amniotic ectoderm;
CH. ALL. MEMB., chorio-allantoic membrane; HND. IN. ALL. MEMB., endoderm
of inner allantoic membrane; VES. IN. ALL. MEMB., vessel in the inner allantoic
membrane.

Bef Be @ Ns
END.1N.ALL-MEMB
VES.IN.ALLMEMB.

In one of the four weeks old embryos an area of combined extra-embryonic
membranes is present on either side of the allantoic stalk, each of which is most
conspicuous. In one of these areas the chorio-allantoic membrane is flattened
against the uterine wall, and the cells of the chorionic ectoderm, mesenchyme
and allantoic endoderm are so compressed that the chorio-allantoic membrane,
in section, has the appearance of a narrow band of undifferentiated tissue (Text-
fig. 17). One of the most obvious modifications is the absence of blood vessels
in the chorio-allantoic membrane in this region (Text-fig. 17). The layer of
endoderm cells lining the inner allantoic membrane is pressed closely against
the chorio-allantoic membrane and there is a thin layer of coagulum between them,
which fills what is left of the allantoic vesicle after the junction of its inner and
outer walls, and which resembles that in the omphaloplacental region described
above. The endoderm cells of the inner allantoic wall are moderately large with
large oval nuclei, but have been compressed by the junction of the two membranes,
until they are smaller than those in the regions of non-attachment. The under-
lying mesenchyme is richly vascularized and obviously feeds and drains the
placenta in this region, since blood vessels are absent in the mesenchyme of the
overlying chorio-allantoic membrane. The mesoderm of the amnion is joined
to the mesenchyme and it and the amniotic ectoderm are normal.

On the opposite side of the allantoic stalk, the inner allantoic membrane and
the attached amnion are in process of joining with the chorio-allantoic membrane

BY H. CLAIRE WEEKES. 535

and consequently the tissues concerned here have a different structure (Text-
fig. 18). The chorio-allantoic membrane is practically normal with blood vessels
present in the mesenchyme. The endoderm cells of the inner allantoic membrane
are greatly enlarged, measuring as much as 0:04 mm. in length, and where they
touch the chorio-allantoic membrane, have attacked the substance of its endodermal
lining, one of the endodermal nuclei having been torn away from its surrounding
cytoplasm on the separation of the outer and inner allantoic walls in the
preparation of the material for sectioning. The cells have square free margins with
large vacuoles at their bases, and on the whole resemble the endoderm cells
described above as lining the inner allantoic membrane in its normal position
and condition. There are as yet no large blood vessels in the mesenchyme under-
lying the endoderm cells, but small vessels are present. The ectoderm cells of
the amnion are slightly enlarged and rounded (PI. xl, fig. 7).

The folds of tissue which surround the blood vessels in their passage across
the allantoic cavity have the same general structure as the rest of the allantoic
tissue. The endoderm cells forming the outer covering of the bridges are
enlarged and many layers deep, and the connective tissue is thick and muscular,
carrying branch vessels as well as the main vessels across the cavity.

UT.

=e

WALL MAT,CAP. EPI. NUC.-

CH.ECT. IN. ALL-END.

AMN. ECT.

Text-fig. 18. Section of the second area of combined foetal membranes in
the allantoplacental region showing the junction of the endodermal lining of the
inner allantoic membrane with the endoderm of the chorio-allantoic membrane;
x 485. AMN. ECT., amniotic ectoderm; CH. ECT., chorionic ectoderm;
EPI. NUC., epithelial nuclei; IN. ALL. END., inner allantoic endoderm;
MAT. CAP., maternal capillary; UT. WALL., uterine wall.

536 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

The apposition of inner and outer allantoic walls results in the formation of”
a short cut for the maternal secretion in its passage to the embryo. Instead of
being collected by the vessels in the chorio-allantoic membrane and passed into
the main right vein and thence across the cavity of the allantoic vesicle into
the allantoic stalk, the secretion passes through the chorio-allantoic membrane
into the vessels of the inner allantois, and thence direct to the allantoic stalk
and so into the embryo. As far as can be ascertained there is no mechanical
pressure upon the inner allantoic membrane which might have effected its union
with the outer allantoic membrane, the embryo lying apart from the region
of junction, but there is no positive evidence that the union has been caused by an
urge for quicker transport of blood to the embryo. In its downward growth the
inner wall of the allantois meets the amnion and the union of the two membranes
is the natural result. This is established before the junction of the inner and
outer allantoic membranes and so the amnion is carried to the placenta by the
agency of the inner allantoic membrane, and obviously can have no new function
in the placental region.

The anticipation of the extension of the allantoic vesicle by the uterine wall
overlying the yolk-sac is as marked as in earlier stages. However, during the
last two weeks of the gestation period when the yolk-sac is rapidly absorbed and
reduced to a small bag, the allantois advances rapidly and the changes which
take place in the uterus are most marked and will be described at Stage C.

UT EPIC. ory MEMB.

ENL.CH.C. FOET. CAP.

Text-fig. 19. Section of the folded allantoplacental region present in one
of the embryos examined; x 450. ENL. CH. C., enlarged chorionic ectoderm
cell; FOET. CAP., foetal capillary; MAT. CAP., maternal capillary; SH. MEMB.,
shell membrane; UT. EPI. C., uterine epithelial cell; UT. GL., uterine gland.

BY H. CLAIRE WEEKES. 537

In one of the embryos examined the allantoplacenta differs from the normal
condition described above. That the variation is individual and not repre-
sentative of a stage in the development of the placenta is shown by its presence
in the one embryo and not in the other embryos of the same female nor in
embryos of females collected at the same time, and hence approximately at the
same stage in development. The embryo is about six weeks old and the allanto-
placenta presents an interesting variation in the form of slight folding of the
maternal and foetal tissues in the region of the main vein and artery, the
folded area being 5 mm. in length and 3:5 mm. in width. The rest of the
placental area is smooth as in all other embryos examined. The folds are not
nearly as definite as the villous folds in the allantoplacental region of
C. tridactylus and L. entrecasteauxzi and in the omphaloplacental region of
T. scincoides, and were not visible to the unaided eye. The degree of folding is
shown (Text-fig. 19; Pl. xxxix, fig. 5), where the foetal tissue is seen fitting
loosely into the shallow maternal crypts. The uterine wall is much thicker than
in the non-folded region. The structure of the mucosa and glands is normal, but
the epithelium and capillaries merit special attention. Although the epithelium of
the folds has not degenerated, the cells are not as uniform in shape as in the non-
pregnant condition, being crowded together in some places until they are cone
shaped (Text-fig. 19). Hach is slightly enlarged, has a rounded free margin and a
central round or oval nucleus. The cells appear to be secretory, being non-
vacuolated, and somewhat resemble those lining the villous folds of
L. entrecasteauxi. In addition the epithelium is not interfered with to any
extent by the maternal capillaries, only a few having penetrated into its substance,
the remainder lying in the mucosa beneath the epithelium. However the whole
area is not as well vascularized as the non-folded region.

Between the maternal and foetal tissues there is a thin remnant of shell
membrane extending over the whole of the placental area. The chorionic ectoderm
cells have not degenerated to the same extent as they have in the non-folded region
and some are larger here than in any other of the allantoplacentae examined, but
the majority are normal in size and have no dividing walls or definite shape. Each
cell contains a large oval or round deeply staining nucleus, which has one or two
nucleoli, and which in some cases almost completely fills the cell. The foetal
capillaries are numerous, the connective tissue is comparatively thick, and the
structure of the endoderm corresponds to that in a normal placenta.

In the non-folded region the maternal capillaries are numerous and have
reached the surface of the epithelium which for considerable stretches has
completely degenerated. The allantoic capillaries have also reached the surface
of the foetal tissue, the chorionic ectoderm cells being few and scattered.

In the allantoplacenta of each of the embryos collected during the last month
of the gestation period the main differences from the placenta described as typically
mature are (a) an increase and enlargement of the maternal and foetal capillaries
to meet the increasing demands of the rapidly growing embryo; (0) the further
degeneration of the uterine epithelium between the maternal capillaries; (c)
pronounced bulging of the maternal and foetal capillaries over the placental face,
the capillaries being closely apposed often with no maternal or foetal cytoplasm
between them, and with their own walls reduced in thickness.

Stage C. The placentae immediately prior to the birth of the foetus.
Females were collected from Barrington Tops, the Blue Mountains and Kiama
(coast) which contained young ranging from within two weeks to one day of

538 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

birth. Altogether four females were collected from Barrington Tops during the
first two weeks of February, 1925, and the young are estimated to be about ten
weeks old. Females were collected from the Blue Mountains during the last two
weeks of January, 1927, and of these, five were kept until the 12th February, when
the young were still unborn although those in the bush were born by the last
week in January. By the 12th February all but two of the females had died, so it
was thought advisable to open the remaining two and fix the contained embryos,
which are taken as being at the stage immediately prior to birth, since it was
obviously the captivity. of the parents and not the condition of the young which
inhibited birth. Three females were collected from Kiama during the first week
of January, 1926, and were kept alive in Sydney until the end of the second week
when the young were born.

The blastocyst taken from a female of the mountain type when the foetus
is within one week of birth measures on an average 2 cm. in length and that
from a coastal type 2:7 cm. When a female is opened up along the ventral body
wall the yolk-sacs of the contained blastocysts are no longer conspicuous as in
earlier stages, each being reduced to a small bag of yolk about 3 mm. in diameter,
which may be in its normal position pressed against the overlying uterus, but is
usually withdrawn from the latter until it lies hidden among the limbs and tail
coils of the embryo. The embryo can be distinctly seen lying among its membranes,
which are thin and greatly expanded by the large amount of embryonic excretion
present in the allantoic vesicle. The excretion oozes out as a clear colourless
fluid when the allantois is punctured. The embryos are well formed with
definite scale markings and when touched wriggle about within their membranes.
The limbs of some of the young lizards within a week of hatching were pressed
against and almost embedded in the yolk-sac and in one instance the tail was
coiled round it several times. This position of the limbs may indicate an effort
on the part of the embryo to aid the withdrawal of the yolk-sac into its body, since
some of the embryos when only about three weeks old were capable of movement
when stimulated and even crawled for some distance when freed from their
membranes.

From the sixth to the tenth week of the gestation period there is no marked
decrease in the size of the yolk-sac, but during the eleventh and last weeks the
reduction is pronounced, the sac decreasing to about one-twelfth its size at six
weeks. The large cells described at Stage B as lining the upper surface of the
yolk-sac now completely cover the sac, and by intense proliferation have filled
the interior, which in section appears to be divided into numerous rounded areas
usually composed of three or four cells surrounding a central blood vessel
(Text-fig. 20, A and B). The cells are efficient yolk absorbers and rapidly empty
the yolk-sac, passing the yolky material to the blood vessels. They are relatively
enormous, each being definitely bounded and containing one large, deeply staining,
irregular nucleus and many small or large vacuoles. The cell cytoplasm is stained
a deep pink by eosin and is obviously packed with yolk material having the same
appearance as the yolk granules. Many of the cells are actively ingesting yolk
granules and their cytoplasm is packed with them, some completely filling the cell,
others being extremely small and obviously prepared by the cells for their passage
into the blood vessels (Text-fig. 20, C). The blood vessels encircled by the
groups of cells are in many cases newly formed, haematopoiesis in the yolk-sac
being active at this stage.

co
it~)

BY H. CLAIRE WEEKES. Be

ALL. MEMB.

ALL.MEMB. C

(FH) (i
/
A, Os {Oe @e

D GHAEGIE

Y.S. END-

Text-fig. 20. A, section of yolk-sac of an eleven weeks old embryo showing
the junction of the allantoic membranes at the base of the sac, the shell membrane
and the “growth” of allantoic tissue; x 26. B, transverse section through the
enlarged endoderm cells in the yolk-saec surrounding a blood vessel; x 562.
C, section of a portion of the yolk-sac wall; x 562. D, section of the membrane
formed by the junction of the outer allantoic membrane with the layers of
somatic mesoderm, yolk-sac endoderm and chorionic ectoderm; x 562.
ALL. MEMB., allantoic membrane; CH. ECT., chorion ectoderm; ENL. END. C.,
enlarged endoderm cell; F. GR. CH. ECT., finger-like growth of chorionic
ectoderm; GR.T., growth of tissue; SH. MEMB., shell membrane; Y. GR., yolk-
granules; Y.S., yolk-sac; Y.S. END., yolk-sac endoderm

540 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

POS. Y.S. ST.

UMB. V.

BY H. CLAIRE WEEKES. 541

With the reduction in size of the yolk-sac space is provided for the further
extension of the allantois and at this stage the allantoic membranes completely
envelop the yolk-sac, meeting and fusing at its base. The fusion is accompanied
by a considerable growth of tissue (allantoic mesoderm and endoderm, 'Text-
fig. 20, A; Pl. xl, fig. 6), which was observed in all the embryos examined and in
the living condition was visible as a circular, flat, white area beneath the yolk-sac,
or in the former position of the yolk-sac when the latter was no longer present.
This “growth” of tissue varies in size, measuring on an average about half the
diameter of the sac in the mountain type of blastocyst. The main mass of tissue
is muscular with blood vessels embedded, and contains numerous yolk spheres
and small dark granules which are possibly the remains of degenerating nuclei.
Here, the vessels from the inner allantoic membrane pass to the outer allantoic
membrane slung in thick folds of mesenchyme (Text-fig. 21, D) and lining the
allantoic cavity the endoderm cells are similar in structure to those described
above (Stage B, Text-fig. 16, A), having narrow bases and swollen apices. In this
region they contain many small and a few large yolk-granules which, together
with those in the main mass of tissue, were possibly derived from the outer
layer of yolk-sac endoderm attached to the chorion and now overlying the outer
allantoic membrane, these granules being passed intact from cell to cell. The
under surface of the mass of proliferated tissue is bound by the chorionic ectoderm
which formed part of the original foetal portion of the omphaloplacenta. The
small yolk-sac now passes out from the encircling folds as indicated in Text-fig.
21, A and B, and into the gut of the embryo, where it is absorbed. The vitelline
artery and vein with their branches are naturally withdrawn and absorbed with
the sac. The yolk-sac stalk is torn away from the allantoic stalk and its position
is marked by a thick proliferation of mesenchyme (Text-fig. 21, C).

The allantoic stalk contains a main artery and vein on either side as in
earlier stages but the vessels are smaller. With the withdrawal of the yolk-sac
and its stalk the allantoic stalk now appears to divide longitudinally a short
distance from the body of the embryo, the upper stalk passing upwards and
carrying an umbilical artery and vein which pass across the allantoic cavity as
described at Stage B, the lower stalk passing downwards and carrying the
remaining umbilical artery and vein directly to the allantoic membrane. In some
embryos obtained from the Blue Mountains on 20th January, 1927, the yolk-sac
was just entering the body and the ventral wall of the embryo was distended
round the sac. After the removal of the sac the allantoic folds which surrounded
it meet and join, a further proliferation of tissue occurring at the junction
(Text-fig. 21, D), which is an extension of the original “growth” of tissue formed
beneath the yolk-sac, and described above. There is a cavity at its centre
containing corpuscles, debris and loose cells which were torn away from the yolk-
sac when removed from the vicinity of the membranes and most of which are

Text-fig. 21. A, diagrammatic representation of the yolk-sac of an eleven
weeks embryo encircled by the allantois. B, diagrammatic representation of the
allantoic membranes immediately after the withdrawal of the sac. C, section of
allantoic stalk showing the proliferated tissue in the former position of the yolk-
sac stalk; x 30. D, section of the mass of proliferated allantoic tissue filling
the former ‘position of the yolk-sac; x 30. ALL. END., allantoic endoderm;
ALL. ST., allantoic stalk; CAV., cavity; CH. ECT., chorion ectoderm; C. DEB.,
cell debris; GR. ALL. T., growth of allantoic tissue; L. UMB. VES., left umbilical
vessels; MUS. T., muscular tissue; POS. Y.S. ST., position of yolk-sac stalk;

SQ. T., squamose tissue; UMB. A., umbilical artery; UMB. V., umbilical vein;
Y.S., yolk-sac.

542 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

now degenerating. The tissue is composed of a thick, inner muscular coat formed
from the allantoic mesoderm, and a thick coat of uniform squamose cells formed
from the allantoic endoderm, the outer layer of which consists of enlarged cells
containing yolk granules. The amnion may be joined to the inner allantoic
membrane and when so joined shares the fate of the allantois at the birth of the
foetus. It is not known what becomes of the allantois at birth, since the young
were born from the females in captivity while unobserved, but there were no
membranes left in the uteri of these females, which were opened immediately upon
the discovery of the birth of the young, which was approximately twenty-four hours
later. It is possible that the young are born with these membranes attached
as described for 7. scincoides (Harrison, 1926), the embryos of which dispose
of their membranes by biting the allantoic stalk and eating them. At birth the
young lizards are quite able to fend for themselves, those born escaping out of
the box and through several rooms into the open where they were found the day
after birth.

The Omphaloplacenta.—As the yolk-sac is reduced in size the area of
omphaloplacentation is naturally correspondingly reduced until, finally, when the
yolk-sac is withdrawn from its position underlying the uterus and the allantoic
membranes have extended under the sac, there is no longer an area of omphalo-
placentation, since one of its main requisites, the foetal blood supply in the yolk-
sac, is being removed with the sac, and its chorionic ectoderm is now part of the
newly extended chorio-allantoige membrane. Allantoplacental modifications of
maternal and foetal tissues extend under the yolk-sac in the place of the former
omphaloplacental modifications, so that the maternal and foetal tissues in this
region are twice modified for placentation. However the position of the omphalo-
placenta is marked until the time of birth by the remains of enlarged chorionic
ectoderm cells which line the mass of proliferated tissue at the junction of the
allantoic membranes beneath the yolk-sac (Text-fig. 21, D).

The Allantoplacenta.—As the allantois grows under the yolk-sac the outer
allantoic membrane unites with the layers of somatic mesoderm and chorionic
ectoderm so that a peculiar arrangement results, where instead of the normal
chorio-allantoic membrane forming the foetal portion of the allantoplacenta, a
membrane is established which contains in addition a layer of yolk-sac endoderm
between the chorionic ectoderm and the outer allantoic membrane (Text-fig. 20, D).
The condition of the foetal tissues of the allantoplacenta is further complicated
by the previous modification of the chorionic ectoderm for its functioning as part
of the omphaloplacenta. However it is remodified for allantoplacentation by its
degeneration over the surface of the placental face, with the exception of the area
underlying the mass of tissue formed at the junction of the allantoic membranes.
Here the chorionic ectoderm is composed of many cell layers, the cells resembling
those of the omphaloplacental region described at Stage B, being enlarged and
irregularly shaped, with the lower layers squamose. The “finger-like’’ down-
growths of cells are present among the folds of the shell membrane and are
sometimes disconnected from the rest of the chorionic ectoderm and lie loose among
the folds. Over the rest of the newly extended allantoplacental area with its
well vascularized outer allantoic membrane, the chorionic ectoderm partly
degenerates (Text-fig. 22, B), and is only represented by the isolated enlarged
cells which serve to attach the foetal tissue to the uterus and which are
characteristic of the main allantoplacental area. The layer of yolk-sac endoderm
beneath the chorionic ectoderm also degenerates, the contained yolk granules being

BY H. CLAIRE WEEKES. 543

absorbed by the underlying allantoic tissue. The degeneration of the chorionic
ectoderm and yolk-sac endoderm is probably directly due to the presence of the
underlying allantoic capillaries, since, in the region of the junction of the allantoic
membranes where the thick pad of tissue underlies the chorionic ectoderm and
yolk-sac endoderm instead of numerous allantoic capillaries, the chorionic ectoderm
and yolk-sac endoderm remain practically intact.

The changes which occur in the allantoic membrane in this region correspond
to those which occur in the main allantoplacental region, the capillaries invading
the overlying degenerating foetal tissue. The numerous capillaries are typically
round, packed with corpuscles and have thin walls, and before the degeneration
of the chorionic ectoderm and yolk-sac endoderm lie embedded in a thick coat of
mesenchyme (Text-fig. 20, D), but after the degeneration of these tissues they
may take up the whole width of the allantois. The pad of shell membrane is in
the same position and in much the same condition as in earlier stages, having the
same coagulum among its folds. The fate of the shell membrane at the birth of
the foetus is not known, it being either absorbed before birth by the foetal
tissues, removed with the foetus at birth or left behind in the uterus and
subsequently absorbed. It is unlikely that the foetal tissues can absorb such a
quantity of membrane in the few remaining days before birth, since it remained
practically intact at the base of the yolk-sac for two months.

The extension of the allantois and the modification of the foetal tissue is
accompanied by a change in the structure of the overlying uterine wall, the
maternal tissue being modified for some distance beyond its proximity to the
allantois, as at Stage A. The uterus passes from a condition of modification for
omphaloplacentation, where it is thick, with small capillaries overlying a layer of
fairly deep columnar epithelium, to a condition of modification for allanto-
placentation, where the epithelium degenerates and the capillaries enlarge and
pass to the surface of the uterus (Text-fig. 22, B). Over some areas the epithelium
may remain intact but in a reduced condition as a layer of small cells, however,
there are large areas where the degeneration is as marked as in the main
allantoplacental region.

Where the uterus was previously slightly folded in the omphaloplacental
region, an interesting condition results when the tissues are remodified for allanto-
placentation (Text-fig. 22, A). Hach fold is mainly filled by vacuolated glandular
tissue, containing many small round nuclei. Between the folds there may be
large oval capillaries while numerous smaller, typically rounded capillaries lie
at the surface of the folds either exposed or covered by a thin layer of epithelium.
In the crevices between the folds where the maternal capillaries are comparatively
widely separated from the foetal capillaries the epithelial cells do not degenerate
but become enlarged and glandular, with narrow bases and swollen apices
(Text-fig. 22, A), and function for food secretion. Early in this investigation
only advanced stages in the development of the placenta were available and upon
the discovery of such an area of placentation it was at first thought to represent
the type of allantoplacentation for L. quoyi. However the study of earlier
stages has clearly shown that this area was originally modified for omphalo-
placentation and its present folded structure is a result of such modification.

The females kept in cavity until the 12th February contained embryos which
are taken to be at a stage immediately prior to birth, where the yolk-sac is
completely withdrawn into the body of the embryo and the allantoic membranes

from both sides of the yolk-sac have met and fused. In the main allantoplacental
P

544

PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

B GL. D.UT EPI.

ENE GEEGInC:

FOET. CAP.

UT.« CH-ALL. SEP

Text-fig. 22. <A, section of the folded uterine wall remodified for allanto-
placentation; x 460. B, section of the allantoplacental region in the former
position of the omphaloplacental region; x 613. D. UT. EPI., degenerating
uterine epithelium; ENL. CH. ECT. C., enlarged chorionic ectoderm cell;
FOET. CAP., foetal capillary; GL, gland; MAT. CAP., maternal capillary ;
SEC. C., secretory cells; UT. EPI., uterine epithelium; UT. & CH. ALL. SHP.,
uterus and chorio-allantoic membrane separated.

BY H. CLAIRE WEEKES. 545

region of these embryos the maternal and foetal tissues are much thinner, the
muscle layers and glands of the uterus being markedly flattened. The maternal
and foetal capillaries are not as numerous nor as full of corpuscles as in earlier
stages and the foetal tissue is easily separated from the maternal wall when
dissecting or sectioning the material.

It is thought that the structure of the uterus changes in anticipation of
allantoplacentation and not as a result of stimulus from the immediate proximity
of the allantois, since it will be recalled that in embryos examined one week after
fertilization, the uterus shows early indications of placental modification when the
allantois is merely a small swelling at the posterior end of the embryo; also the
uterus overlying the edge of the yolk-sac is modified for placentation before the
extension of the allantois to its vicinity; in addition, in some embryos eleven
weeks old the allantois underlying the yolk-sac is covered by a thick coat of
chorionic ectoderm cells and the conditions resemble those of omphaloplacentation,
the uterine wall being in contact with and having its materials absorbed by the
specialized chorionic ectoderm, yet here the uterus does not retain its omphalo-
placental modifications, but is well modified for allantoplacentation; also the uterus
is modified for allantoplacentation even when separated from the allantois by a
thick pad of shell membrane. However it may be regarded as peculiar that the
modification progresses roughly parallel with the extension of the allantois and
does not occur simultaneously over the uterus, and this may be taken as indicating
a direct relationship between the two processes, but it is most probably a result
of the uterus retaining its omphaloplacental modifications and only replacing
them by allantoplacental modifications as the foetal omphaloplacentation yields to
allantoplacentation. .

The modification of the foetal and maternal tissues for allantoplacentation as
late as the last two weeks of the gestation period shows that the allantoplacenta
is essential until the birth of the foetus.

4. Period after birth.

Females were available in a condition ranging from one day to one week
after the birth of the young. In these females the uteri were thick, much
convoluted and distinctly restricted into incubatory chambers. The wall of the
uterus extracted from one of the females one day after the birth of the young
measures about 0-4 mm. in its thickest part, the muscle layers being thicker than
in the normal non-pregnant condition and the glands having partly regained their
normal shape and structure. The cytoplasm of the gland cells encloses small
yolk granules derived from the debris in the cavity of the uterus, and the granules
are also present in the epithelial cytoplasm. The inner surface of the uterus is
“thrown into deep folds which are covered by a thin layer of regenerating
epithelium. However the capillaries are still concentrated at the surface of the
mucosa and in places are almost exposed and have the typical rounded appearance
of the capillaries in the allantoplacental region. The epithelial cells are largest
in the crevices between the folds, being narrow and columnar with an oval
nucleus at the base. The uterine cavity contains a dark staining coagulum mixed
with yolk spheres and cell debris which fills the crevices between the folds and is
being rapidly absorbed by the uterus.

At the end of the first week after the birth of the young the uterus is more
normal in structure, but is still obviously influenced by the preceding gestation
period. Its wall is thinner but the glands have not completely regained their

546 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

normal structure, the nuclei being still grouped closely together. The mucosa .
contains many more small capillaries than in the normal condition, but the
epithelial cells, although as yet comparatively small, are uniformly arranged over
the surface of the uterus, the majority being definite and columnar. Most of

the coagulum in the cavity has been absorbed and there are fewer yolk spheres
in the mucosa.

IV. CoMPARISON WITH THE LizarpDs Chalcides tridactylus, Lygosoma (Liolepisma)
entrecasteauxi AND Tiliqua scincoides AND WITH THE MARSUPIAL Perameles.

(a). Comparison with C. tridactylus and L. entrecasteauzi.

The structure of the allantoplacenta in L. entrecasteauxi is so remarkably
similar to that in C. tridactylus that it is unnecessary to compare and contrast
the placenta in each with that in LZ. quoyi. L. entrecasteauxi has been chosen in
preference to C. tridactylus for a detailed comparison with L. quoyi, since the
author has described the placentation of that lizard in conjunction with Professor
Harrison (Harrison and Weekes, 1925), and since material is available for
examination in the sectioned and unsectioned condition.

The discovery of allantoplacentation in L. entrecasteauzi and in L. quoyi,
two species of the one genus Lygosoma, is more important than might be at first
supposed, since their methods of allantoplacentation differ more than might be
expected after a study of the placenta in related species of mammals. The two
lizards were collected from Barrington Tops and Mount Kosciusko, although
LL. quoyi was restricted to the marshy regions and L. entrecasteauzi to the hills
surrounding them. JL. entrecasteauxi is a much smaller lizard than L. quoyi,
being about one-third the size of the coastal type of the latter, and its eggs are
correspondingly smaller. However, although the yolk-sac is only about one-third
the size of that in ZL. quoyi the omphaloplacenta is more highly developed. In
the stages available for examination there was no indication of the abnormal
growth of the extra-embryonic mesoderm into yolk-sac, but this may be evident in
earlier stages. The omphaloplacenta of L. entrecasteauxi differs from that in
L. quoyi in that the cells of the uterine epithelium are much larger and more
uniform in structure; the chorionic ectoderm cells are also larger, there being a
single row of evenly enlarged cells with large dark nuclei, in the place of the
multilayered chorionic ectoderm in ZL. quoyi with its cells varying in length and
shape; the area of omphaloplacentation is relatively greater and this is thought to
be due to the absence of the thick pad of shell membrane in the embryos of
L. entrecasteauzi examined and its invariable presence in the omphaloplacental
region of L. quoyi.

Harrison and Weekes (1925, p. 475) describe the maternal portion of the
allantoplacenta as being restricted to “a fusiform to elliptical opaque whitish area’”’
and the extent of the foetal placentation as slightly exceeding that of the
maternal placental area. In L. quoyi, however, the placental area is not restricted,
modifications of maternal and foetal tissue occurring over the entire area
embraced by the allantois. In L. entrecasteauxi the elliptical placental area is
marked by complicated villous folds of the maternal wall which can be seen with
the unaided eye, but in L. quoyi the placental face is smooth and discernible only
in sectioned material. As well as the differences in general appearance, the
histological differences between the two allantoplacentae are marked. In the
placenta of L. entrecasteauzxi two elliptical epithelial sheets with every cell
clearly bounded are closely apposed but are in no way joined or fused (Text-

BY H. CLAIRE WEEKES. 5AT

fig. 23). The epithelium lining the maternal ridges is composed of enlarged ciliated
cells. The villous folds impress their form upon the closely underlying ciliated
chorionic ectoderm causing irregularities in its surface and variations in the
size of the cells, some growing up into the crypts and being extremely elongated,
in all cases being twice as long as they are wide. The maternal folds are fed
by large capillaries while the foetal vascular network lies close at the base of the
chorionic cells and is bound internally by a sheet of mesenchymatous connective
tissue of varying thickness. In L. quoyi the maternal wall is not covered by
enlarged ciliated epithelial cells, there being at the most a thin bounding layer
of epithelial protoplasm with occasional nuclei embedded; there is a network of
small rounded maternal capillaries instead of the few large capillaries in the
villous folds of L. entrecasteauxi, and these are exposed at the surface of the
uterus and do not lie beneath a thick epithelial layer; the foetal tissue is attached
to the maternal wall, the attachment being efficient although superficial; the cells
of the chorionic ectoderm do not become enlarged but degenerate, allowing the
allantoic capillaries to bulge at the surface, so that the maternal and foetal
blood streams are in close apposition.

—— 62

~ Ch.Ect. AU. Cap.

Text-fig. 23. Section of the elliptical allantoplacental area in
L. entrecasteauci. All. Cap., allantoic capillary; All. End., allantoic
endoderm; Ch. Ect., chorionic ectoderm; Ut. Cap., uterine capillary ;
Ut. Epi., uterine epithelium; Ut. Muc., uterine mucosa.

Hence it can be seen that the allantoplacenta in ZL. quoyi is fundamentally
different from the elliptical and _ restricted allantoplacental region in
L. entrecasteauzi, since food transition in L. quoyi is carried on by the apposed
maternal and foetal bloodstreams and not by the glandular activity of the uterine
epithelium and the phagocytic and absorbing powers of the chorionic ectoderm as
in L. entrecasteauzxi.

The omphaloplacenta in C. tridactylus (Giacomini, 1891, p. 348) “se forme
tardivement et reste rudimentaire” consisting of enlarged chorionic ectoderm cells
in the region of the yolk-sac. No mention is made of any modification of the uterine
tissues in this region. It is possible that the development of the omphaloplacenta

548 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

may be restricted by the small size of the eggs (2:5 to 3 mm.) in such a.
comparatively large lizard. The growth of the extra-embryonic mesoderm into
the yolk-sac in L. quoyi has no parallel in C. tridactylus since Giacomini says,
1891, p. 3438, “tant que les oeufs ne sont pas arrivés 4 un certain degré de
développement, la connexion immédiate entre l’ectoblaste et l’entoblaste vitellin
persiste telle qu’elle a été décrite. Le feuillet moyen s’arréte aux limites de la
connexion, de sorte que, pendant quelque temps, la paroi vitelline, sur ce point,
est privée de sa structure caractéristique, et ne se trouve pas vascularisée. Dans
des états avancés de développement, le mésoderme envahit aussi la région en
question, laquelle, alors, est également vascularisée’”. This is the only reference
Giacomini makes to the growth of the extra-embryonic mesoderm over the yolk-
sac and it is taken to mean that the sinus terminalis remains quiescent for some
time at the outer limits of the area of junction of the chorionic ectoderm with
the yolk-sac endoderm, finally continuing its growth round the sac in the normal
way between the ectoderm and endoderm.

An interesting and important phenomenon has been observed in the placentation
of L. entrecasteauxi since the publication of the papers by Harrison and Weekes,
1925, and Weekes, 1927. The area of apposed maternal and foetal tissue in
L. entrecasteauxi other than the omphaloplacental and the elliptical allantoplacental
areas was regarded as unimportant and non-placental in the investigation in
1925, the maternal and foetal tissues being so thin and flattened here that it was
difficult to make a histological study of them. However in one of the series of
sections a relatively thick area was recently observed, attention being first arrested
by the very close attachment of the foetal to the maternal tissues in this region
which persisted even when the tissues in the elliptical allantoplacental area
were fairly widely separated. Upon a closer examination it was found that the
structure of this region corresponds closely to that of the allantoplacenta in
L. quoyi. The maternal and foetal epithelial cells have degenerated, the epithelial
nuclei in the maternal wall are grouped between the capillaries and have the
same typical rounded appearance as in L. quoyi, and the foetal tissue is attached to
the maternal tissue by scattered enlarged chorionic ectoderm cells. The only
obvious differences between the structure of this region and that of the allanto-
placenta in L. quoyi are the smaller number of maternal and foetal capillaries,
their ellipsoid cross section and the presence of a thick coat of mesenchyme
underlying the foetal capillaries. It is thought that in younger stages in the
development of L. entrecasteauxi this specialized region may be more extensive.

It will be recalled that at Stage B a condition of allantoplacentation is
described for one of the embryos of L. quoyi examined, where the uterine wall
and chorio-allantoic membrane are thrown into rudimentary folds and where the
epithelium of the maternal folds has not degenerated, and where, in addition, the
chorionic ectoderm cells have not degenerated to the same extent as in the
allantoplacenta of other embryos, a few of them being larger than in any of
the other allantoplacentae examined. When this folded area is compared with
the elliptical placental area of L. entrecasteauxi the resemblances are noticeable.
The position of the folds in the region of the main longitudinal artery and vein
resembles the position of the folds in L. entrecasteauxi; the maternal epithelial
tissue obviously plays the important part in food secretion since the capillaries
are not present at the surface of the uterus, and since the epithelial cells are
enlarged and obviously modified for secretion; the chorionic ectoderm cells are
comparatively large and contain enormous nuclei, the shape of the cells being to

BY H. CLAIRE WEEKES. 549

some extent modified by the shape of the overlying uterine wall; and finally,
the foetal capillaries lie at the bases of the chorionic ectoderm cells.

There are, however, many differences. The area in L. quoyi, although in a
much larger lizard, is only half the width of that in L. entrecasteauxi and the
folds are only rudimentary when compared with the well formed complicated folds
in that lizard; the uterine epithelial cells vary in shape and size and are not
ciliated nor uniformly enlarged nor arranged; the chorionic ectoderm cells are
neither ciliated nor granular and are not definite in shape, cell boundaries being
recognizable only at the free margins and in addition in some places they are
degenerating and the allantoic capillaries passing to the surface; there is no
deep layer of mesenchymatous tissue underlying the chorion and the allantoic
capillaries.

The significance of the occurrence of the second type of allantoplacentation in
i. entrecasteauzi and of the folded area in one of the embryos of L. quoyi examined
will be discussed below.

(bo). Comparison with Tiliqua scincoides.

T. scincoides is a much larger lizard than either L. entrecasteauzi or
L. quoyi and is slightly larger than C. tridactylus. The habitat of this lizard
resembles that of L. quoyi in that it varies from sea level to an altitude of 3,000
feet. The pregnant females contain a greater number of young than those of
L. quoyi, the two described by Flynn (1923) containing eleven and fifteen. The
general disposition of the blastocyst is the same as in L. quoyi.

According to Flynn the yolk-sac circulation of TJ. scincoides extends over
the outer surface of the sac with the exception of a small area at the lower pole.
In L. quoyi the circulation completely surrounds the sac. Flynn notes (1923,
p. 75) that ‘“‘a sinus terminalis could not be definitely made out”. This was since
discovered (Weekes, 1927) to be due to the downgrowth of the extra-embryonic
mesoderm into the yolk-sac as in L. quoyi. The omphaloplacenta in T. scincoides
(Weekes, 1927) is much more highly specialized than that of L. quoyi, the uterine
wall being thrown into definite villous folds which are visible to the unaided
eye, and the cells of the epithelium are much larger and more regular than those
of the maternal epithelium in L. quoyi. It is possible that the rudimentary
folding of the uterine wall in L. quoyi is the forerunner of such a specialized
condition. The most outstanding differences between the omphaloplacentae are
the regular folding of the foetal tissue in J. scincoides to fit into the maternal
erypts and the regular modification of the chorionic ectoderm cells.

The omphaloplacenta in both lizards is of the same type. the food transition
being carried on by apposed maternal and foetal epithelial faces, and the foetal
tissues vascularized by vessels embedded in the yolk-sac and separated from it by
the extra-embryonic coelome.

The general relationship between the allantois and the embryo is the same
in both lizards with one exception. In TViliqua (Flynn, 1923, p. 77) the allantoic
stalk passes downwards and outwards while in L. quoyi it passes upwards and
outwards. The method of transference of blood vessels across the allantoic
cavity in the two lizards has been compared at Stage B. In both lizards the
inner allantoic membrane fuses with the amnion, but as Flynn does not mention
any connection between the chorio-allantoic and the inner allantoic membranes. it
seems that as far as those lizards for which placentation has been described are
concerned, this arrangement is peculiar to L. quoyi. The area of allantoplacenta-

550 PLACENTATION IN LYGOSOMA (HINULIA) QUOYi,

tion in TJ. scincoides is smaller than that in L. quoyi since its extension is.
restricted by the central portion of the allantois alone becoming attached to the
chorion, the marginal zone being free and unattached. Flynn gives no figures
illustrating the histological relations in the allantoplacenta of Tiliqua and it is
therefore difficult to compare the placenta with that in L. quoyi. He says
(19238, p. 76) that “‘the union between chorion and uterine epithelium is very
intimate. The uterine epithelium apparently consists of a single layer of very
flattened cells, while the chorionic ectoderm has proliferated greatly, is much
vacuolated, resembling a typical plasmodium and is formed in the main of
markedly enlarged cells with large nuclei and connected together by amoeboid
processes. These processes insinuate themselves into and between the maternal
cells in much the same way as Hill has described for the chorionic cells in the
formation of the metrioplacenta of Dasyurus viverrinus”. No special mention is
made of the maternal circulation and in describing the foetal circulation Flynn
says (1923, p. 77) that the allantoic vessels “‘ramify through the mesenchymal:
layer of the placental face’, but makes no mention of any unusual position of
the allantoic capillaries. It is evident from these observations and the above
description of the placental area that there is not the same degeneration of
epithelial surfaces and apposition of capillaries as in L. quoyi. The “very flattened
cells” of the uterine epithelium may present a stage intermediate between the
condition of enlarged cells in C. tridactylus and L. entrecasteauxi and of
degenerated epithelium in JL. quoyi. Hence it is deduced that the maternal
and foetal epithelial tissues in Tiliqua, although not enlarged to the same extent
as in C. tridactylus or L. entrecasteauxzi, carry on the function of food transition
and therefore the type of placentation differs essentially from that in L. quoyi.
However, the two types have in common the attachment of the foetal tissues
to the maternal wall, both being of the conjoint type, and the establishment of
this attachment by enlarged chorionic ectoderm cells which do not invade the
maternal tissue to any extent, the attachment in each being superficial.

(c). Comparison with Perameles.

Perameles is the only marsupial for which allantoplacentation has been
recorded. The placentation of Perameles was described by Hill (1897), redescribed
by Flynn (1923), and these authors hold different opinions as to the nature of
the placenta. Hill claims (1897, p. 387) that the uterine mucosa undergoes
hypertrophy; that the vessels in the mucosa increase in size and number; that
the uterine epithelium changes into a vascular syncytium, the nuclei becoming
grouped together in nests situated in lobular projections of the deeper surface
of the syncytium; and that the maternal capillaries pass up between the syncytial
lobules and form a network beneath the epithelial protoplasm. He claims further
that the embryo becomes attached to the prepared maternal wall by means of
enlarged chorionic ectoderm cells, which eventually degenerate over the placental
area proper; that the allantoic capillaries now directly reach the vascular surface
of the maternal placental syncytium to which they become intimately attached,
dipping down into the depressions on its surface and forming a regular inter-
locking system, and that finally the foetal and maternal blood streams are now
only separated by their thin endothelial walls and perhaps a layer of syncytial
protoplasm.

Flynn claims that the chorionic ectoderm does not completely degenerate but
actively invades the maternal tissues and thus brings the type of placentation

BY H. CLAIRE WEEKES. 551

found in Perameles in line with that found among the eutherian mammals, more
especially the Carnivora “where there are the same characteristics of passivity of
uterine epithelium and activity of the trophoblast’”’.

By the kind permission of Professor Harrison of the Zoology Department,
University of Sydney, I have been able to obtain some of the Perameles material
prepared by Hill and used by him and Flynn. This consists of two slides, one
containing sections of the placenta of an embryo 7 mm. in direct length and
the other of one 1-5 mm. in length. An examination of this material has convinced
me that Flynn’s interpretation of the structure of the placenta is the correct one.
The most important and convincing evidence is the invasion of the deep maternal
syncytial lobules by the large nuclei of the trophoblast, which is so plainly evident
that as Flynn himself says it is impossible that there can be any other conception
than the one he suggests. Flynn has ably and thoroughly discussed the placenta
in Perameles (1923), and there is no need for further comment here, but while
I agree with his interpretation of the structure of the placenta I do not support
his claim that its similarity to that of the placenta in some of the Carnivora is of
fundamental phylogenetic importance.

The relationship between the embryo, its yolk-sac and the allantois in L. quoyi
is somewhat similar to that in Perameles, allowance being made for the difference
in the relative size of the yolk-sac and the allantois in each, the yolk-sac in
Perameles occupying the main portion of the blastocyst and the allantois being
restricted to the remaining portion. In each the embryo is sunk down into the
yolk-sac being partially surrounded by the yolk-sac wall and the yolk-sac is
vascularized by one main artery and vein. But the vascularization does not
completely cover the sac in Perameles as in L. quoyi, there being a definite sinus
terminalis present, and the method of the growth of the extra-embryonic mesoderm
over the sac is normal. The vascularization of the allantois is similar in both,
each of the two main arteries being accompanied by one of the veins in its
ramifications over the placental face, the arrangement being confined to the
main trunks. However there are none of the “cellular bridges” passing across
the allantoic cavity in Perameles as in L. quoyi, possibly on account of the small-
ness of the allantoic vesicle and the consequent lack of any need for a short cut
for the vessels. In both, the allantoic stalk enters the body of the embryo
immediately posterior to the yolk-sac stalk, but in L. quoyi both stalks are joined
by mesenchyme and have the appearance of one.

There is little similarity between the omphaloplacentation of these forms,
since in Perameles the uterine epithelium degenerates to form a syncytium with
the maternal capillaries at the surface, the capillaries being closely apposed to a
flat layer of chorionic ectoderm which is separated by blood vessels from a layer
of enlarged yolk-sac endoderm cells.

According to Flynn’s interpretation of its structure, the allantoplacenta in
Perameles consists of a fused area of complexly folded maternal and foetal tissue.
The maternal wall is thick and modified for the attachment of the foetal tissue
by the proliferation of the nuclei of the epithelium and their migration to the
deeper parts of the epithelium which has now markedly thickened, the result
being the formation of a syncytium in which the deeply situated nuclei assume a
particular form and arrangement. Flynn (1923, p. 137) says “the nuclei become
aggregated mainly in rounded masses or nests situated in the lobular projections
of the syncytial protoplasm. The lower surface of the syncytium has a wavy
appearance due to the presence of the lobules . . . at this stage the syncytium

Q

552 PLACENTATION IN LYGOSOMA (HINULIA) QUOYz,

is well vascularized, each capillary being enclosed in its delicate endothelial layer”. ©
The maternal capillaries pass up between the lobules and form a network at the
surface of the epithelial protoplasm. To this prepared maternal tissue the chorion
attaches itself by a single layer of enlarged ectoderm cells. These cells divide
(p. 141) ‘“‘to give rise to nucleated groups in which cell outlines have disappeared.
. . . At various points these nuclei invade the uterine syncytium and the
remaining basal cells of the trophoblast layer form the cytoblast or cytotropho-
blast”. Further (p. 167) “the outward migration of the basal cytoblast cells where
converted into plasmodiblast gives opportunity for the maternal and foetal vessels
to come into intimate apposition . . . all maternal vessels have definite endo-
thelial walls, hypertrophy of the endothelial cells does not occur and lacunae are
not formed’. In my own observations most of the maternal capillaries were seen
to lie immediately above the allantoic capillaries, both sets of capillaries being
round.
The following are the main differences between the placentation of Perameles
and that of L. quoyi.

1. The uterus in Perameles is much thicker than that in LZ. quoyi, and the uterine
epithelium is deeper both before and after it is modified for placentation, and when
modified is constricted into lobules containing distinct nests of nuclei, which arise by
proliferation.

2. There is no shell membrane and the foetal tissue is attached to the maternal
wall before the fusion of the allantois with the chorion.

3. The enlarged chorionic ectoderm cells proliferate and invade the maternal
syncytium, firmly fixing the foetal tissue in position and absorbing food by their
phagocytic action. i

However the points in common are outstanding and fundamental.

1. The uterine epithelium in ZL. quoyi, although a narrow layer degenerates and
forms a syncytium containing the rounded, palely staining nuclei which resemble those
of the syncytial lobules in Perameles in structure and in that they “lack staining
qualities” (Flynn, p. 137).

2. The maternal capillaries multiply and invade the degenerating epithelial cyto-
plasm and pass to the surface of the uterus. f

3. The foetal tissue is attached to the maternal wall by enlarged chorionic
ectoderm cells which remain throughout the life of the placenta.

4. The maternal and foetal capillaries are closely apposed, being separated only by
their thin endothelial walls and perhaps a layer of maternal and foetal cytoplasm.
The capillaries have definite endothelial walls, hypertrophy of the cells does not occur
and the foetal capillaries lie beneath the maternal.

Some of the differences, however, are due to the natural conditions in
L. quoyi. The narrowness of the uterine epithelial layer does not lend itself to
the formation of a deep syncytium, and the presence of the shell membrane hinders
the early fixation of the chorion to the uterine wall. So that, the only important
difference between the placentae is, that in Perameles the chorionic ectoderm
proliferates and invades the maternal syncytium, whereas in L. quoyi, with the
exception of scattered enlarged cells, it degenerates. Hence can be seen the
similarity between the two types of placentation and the reason for the statement
in the previous paper on the placentation of L. quoyi (Weekes, 1927, p. 29) that
the allantoplacenta in L. quoyi more closely resembles that of the Mammalia than
any hitherto recorded in a reptile.

V. THEORETICAL CONSIDERATIONS.

A discussion of the relationship of the placentation of L. quoyi with that of
other reptiles, and of its similarity to the placentation of the marsupial, Perameles,

BY H. CLAIRE WEEKES. 553

must be deferred until an examination of other viviparous forms of reptiles is
made. Also the as yet undescribed placentation of H. whitei and EH. striolata,
which closely resembles that of L. quoyi, must be taken into consideration.
However, attention is called to a statement (Weekes, 1927, p. 29) that “in the
genus Lygosoma two members, namely L. entrecasteauri and L. quoyi have each
developed distinct types of placentation”. This is true for the folded elliptical
area of placentation in L. entrecasteauzxi, but with the discovery of the second
area of placentation (described above) the difference between the two types is no
longer so marked. It may be that the placentation in these two lizards is related,
the common ancestral type being similar to that in L. quoyi, the placentation of
L. entrecasteauxi becoming independently specialized by the development of the
restricted elliptical region; or it may be that placentation was independently
acquired by each, by L. entrecasteauxi before L. quoyi, this supposition being
supported by the absence of shell membrane in L. entrecasteauxzi and by its
presence and the rudimentary condition of the omphaloplacenta in L. quoyi. The
folded area in one of the embryos of ZL. quoyi examined might be regarded as
representing a developing area in LZ. quoyi such as occurs in L. entrecasteauzi,
but it is probably only a freak in placental development, since it occurs in only
one of the many embryos examined.

VI. Summary and Conclusions.

The placentation of the Scincid lizard Lygosoma (Hinulia) quoyi is described
in detail in the present communication and compared and contrasted with that
in the Scincid lizards Chalcides tridactylus, Lygosoma (Liolepisma) entrecasteauxri
and Tiliqua scincoides and with that in the marsupial Perameles. The following
is a Summary of the work and of the main conclusions drawn.

1. The lizard is not strictly viviparous, there being a shell membrane present which
accumulates at the base of the yolk-sac as a thick pad in advanced stages of development.

2. An omphaloplacenta is present which even in its mature condition is less specialized
than that in L. entrecasteauxi, T. scincoides, EH. whitei and E. striolata.

3. An allantoplacenta of the conjoint type is present consisting of degenerated
maternal and foetal epithelial tissue and apposed maternal:and foetal blood streams.

4. A modification of the structure of the allantoplacenta occurs in some embryos by
the junction of the inner allantoic membrane and the amnion with the placental area.

5. The right umbilical artery and vein pass to the allantoplacental region across the
allantoic cavity and the method of the establishment of its passage is described.

6. The placentation of ZL. quoyi is described as being different from that in
L. entrecasteauxzi and C. tridactylus, where the function of food transition is mainly
carried on by the modified maternal and foetal epithelial tissues.

7. The occurrence of a second area of allantoplacentation in ZL. entrecasteauwaxi is
recorded, the structure of which resembles that of the allantoplacentation in L. quoyi.
No deductions are made at this stage, a further study of possible placentae among the
other viviparous members of the genus Lygosoma being necessary.

8. After an examination of some of the Perameles material used by both Hill and
Flynn, Flynn’s interpretation of the structure of the allantoplacenta in Perameles is
supported in this communication.

9. The type of allantoplacentation in L. quoyi is found to be essentially similar
to that in the marsupial Perameles, with differences in detail.

References.

CALDWELL, 1895.—On the Arrangement of the Embryonic Membranes in the Marsupial
Animals. Quart. Journ. Micr. Sci., xxxvi. 3
Duvau, 1892.—Le Placenta des Rongeurs. Journ. de VAnat. et de la Phys.

FLYNN, 1922.—The Phylogenetic Significance of the Marsupial Allantoplacenta. Proc.
Linn. Soc. N.S.W., xlvii, 4.

554 PLACENTATION IN LYGOSOMA (HINULIA) QUOYI,

FLYNN, 1923.—The Yolk-sac and Allantoic Placenta in Perameles. Quart. Journ. -
Micr. Sci., \xvii, 1. ;

, 1923.—On the Occurrence of a true Allantoplacenta of the conjoint type in an
Australian Lizard. Rec. Aust. Mus., xiv.

GIACOMINI, 1891.—Matériaux pour L’étude du développement du Seps chalcides. Arch.
Ital. de Biol., xvi. (Reprint.from Monitore Zool. Ital., ii, Nos. 9-10.)

, 18914.—tiber die Entwickelung von Seps chalcides. Anat. Anz., vi.

, 1893.—Contribution 4 la connaissance des annexes foetales chez les reptiles,
Jer et 2e, Note préventive. Arch. Ital. de Biol., xviii. (Abstract of paper in
Monitore Zool. Ital., iv, No. 7, 1893.)

, 1893.—Sur l’oviducte des Sauropsides. Arch. Ital. de Biol., xxi, 1894. (Abstract
of paper in Monitore Zool. Ital., iv, Nos. 10-12.)

, 1893.—Nouvelle contribution 4 la connaissance plus parfaite des annexes foetales
chez les reptiles; Reception du sac vitellin et de lallantoide dans la cavité abdominale.
Arch. Ital. de Biol., xxi, 1894a. (Abstract from paper in Monitore Zool. Ital., iv,

INOrsiG;, 18:93)

HaackeE, 1885.—Uber ein neue Art uteriner Brutflege bei Reptilien. Zool. Anz., viii.

HARRISON, 1926.—Roy. Zool. Soc. N.S.W., iv, part 5.

HARRISON AND WEEKES, 1925.—On the Occurrence of Placentation in the Scincid lizard
Lygosoma entrecasteauai. Proc. LINN. Soc. N.S.W., 1.

Harper, 1900.—The Sexual Season, Quart. Journ. Micr. Sci., xliv.

Herr, 1924.—Das Corpus Luteum der Zauneidechse (Lacerta agilis). Zeitschr. Mikr.
Anat. Forsch. 1 Band, 1 Heft.

Hitt, 1897.—The Placentation of Perameles. Quart. Journ. Micr. Sci., xl.

HUBRECHT, 1889.—The placentation of Hrinacews europaeus. Quart. Journ. Micr. Sci.,
2OOG,

JENKINSON, 1913.—Vertebrate Embryology.

LILLE, 1908.—The Development of the Chick.

MARSHALL, 1910.—Physiology of Reproduction.

Minot, 1891.—A Theory on the Structure of the Placenta. Anat. Anz., vi.

WEEKES, 1927.—A Note on Reproductive Phenomena in some lizards. Proc. LINN. Soc.
N.S.W., lii- 2. ;

EXPLANATION OF PLATES XXXVIII-XL.
Plate xxxviii.

Fig. 1. Photomicrograph of section of omphaloplacental region of embryo collected
during the second week of the gestation period.

Fig. 2. Photomicrograph of section of yolk-sac of a young embryo showing the down-
growth of mesoderm into the yolk-sac.

Plate xxxix.

(Jt)

Fig. Photomicrograph of section of yolk-sac of six weeks old embryo showing the

dense coagulum with corpuscles and scale-like appearance.

Fig. 4. Photomicrograph of section of mature allantoplacental region showing apposition
of maternal and foetal tissues.

Fig. 5. Photomicrograph of section of folded allantoplacental region present in one of
the embryos examined.

Plate xl.

Fig. 6. Photomicrograph of section of yolk-sac of eleven weeks old embryo showing
the enlarged endoderm cells, growth of allantoic tissue and shell membrane.

Fig. 7. Photomicrograph of section of allantoplacental region showing the junction of
inner allantoic membrane with the chorio-allantoic membrane.

LEGENDS TO PLATES XXXVIII-XL.

A.C., allantoic cavity; AMN., amnion; B.V., blood vessels; C., coagulum; C.E.,
chorionic ectoderm; E. CYT., endodermal cytoplasm; #.C.H.C., enlarged chorionic ectoderm
celi: E.E.C., extra-embryonic coelome; E.N. E.C., enlarged endoderm cell; F.C., foetal
capillary; F.F.T., folded foetal tissue; F.GR., finger-like growth; GR. T., growth of
tissue; I.A.M., inner allantoic membrane; M.C., maternal capillary; P.S.C., peculiar shaped
cells; SP. M., splanchnic mesoderm; SC. M., scale-like material; SO. M., somatic mesoderm ;
S.M., shell membrane; U.E., uterine epithelium; UT., uterus; Y.S., yolk-sac; Y.S.E.,
yolk-sac endoderm.

NOTES ON AUSTRALIAN MARINE ALGAE. v.
By A. H. S. Lucas, M.A., B.Sc.

(Plates xli-xlviii.)
[Read 30th November, 1927.]

Contents,

Algae of Michaelmas Cay.

Chlorophyceae from Bowen.

Two forms of Codium.

Notes on Caulerpa. Edible Chlorophyceae.
Fucoideae. Distribution Notes.

Two new species of Chondria.

Oorow rer

1. Algae collected at Michaelmas Cay, 27 miles off Cairns, Qld., by
T. Iredale and G. P. Whitley in June, 1926.

Under the auspices of the Great Barrier Reef Committee a bore was put down
in Michaelmas Cay, a narrow sandbank lying 27 miles off Cairns. The Cay is about
300 yards long and runs from N.E. to S.W. and is quite low. Rough accommodation
was provided on the Cay for the workmen and naturalists. Messrs. Iredale
and Whitley spent a month there engaged in the study of the fauna of the reefs
and in collecting zoological specimens for the Australian Museum, Sydney. At
my request they kindly undertook to gather the algae as well.

The prevalent winds blow from the S.E. On that side corals flourish, but
algae cannot obtain a foothold on the living coral. On the N.W. are dead corals
in abundance, and the weeds grow freely on the blocks, from sea level to a depth
of some two fathoms at least.

A first and larger collection was unfortunately swept away by an unusually
high and unexpected tide, and lost. A second collection included the following
species:

ee

CHLOROPHYCEAE.
Boodlea coacta (Dickie) Murray. New for Australia.
Dictyosphaeria favulosa (Mert.) Decaisne.
Valonia Forbesiti Harv. New for Australia.
Halimeda opuntia (Linn.) Lamour.
Halimeda tuna (Ellis and Soland). Lamour.
Caulerpa clavifera (Turn.) J. Ag.
PHAEOPHYCEAE.
Hydroclathrus cancellatus Bory.
RHODOPHYCEAE.
Gracilaria taenioides J. Ag.
Hypnea seticulosa J. Ag.
Laurencia obtusa (Huds.) Lamour.
Laurencia rigida J. Ag.
Digenea simplex (Wulf.) J. Ag.
Acanthophora orientalis J. Ag.
Leveillea jungermannioides (Mart. and Hering) Harv.
Ceramium clavulatum (Mont.) J. Ag.
Jania rubens Lamour.

556 NOTES ON AUSTRALIAN MARINE ALGAE, V,

2. Chlorophyceae from Bowen, Port Denison.

Mr. E. H. Rainford, of Bowen, has favoured me with numerous valuable
consignments of algae from that locality. He has very generously put himself to
the labour of freely, carefully and judiciously collecting the seaweeds growing on
the reefs of Stone Island and the shores generally of Port Denison. He has
already sent some 50 species.

Sonder in his Algen des tropischen Australiens, 1871, recorded and described
a number of forms sent to him from Port Denison by Fitzalan and Kilner. Some
of these have been gathered by Mr. Rainford, and also some others not noted by
Sonder. The Chlorophyceae are particularly interesting.

Boodlea coacta (Dickie) Murray was found in Australian waters for the first
time around the Michaelmas Cay, as recorded above. It is evidently abundant
in Port Denison.

Anadyomene Brownii (Gray) J. Ag. This was described from Port Denison
as a new species under the name A. Muelleri, by Sonder, but he had been
anticipated by Gray. Mr. Rainford sent a number of excellent specimens. The
plant also occurs around Celebes.

Dictyosphaeria favulosa (Mert.) Decaisne. A fine suite of specimens. Not
recorded by Sonder, but known from the Sandwich and Friendly Islands and
from Ravak and north Australia. Another species, D. sericea Harv., is found on
the south-west and southern coasts of Australia and around Tasmania. Harvey
thought it might be a form of D. favulosa but gave it a specific name. Agardh
regarded it as distinct.

Valonia confervoides Harv. A single specimen, apparently rolled into a ball
by the waves, so that at first I thought I was dealing with a globular Codium.
It has previously been recorded from Stone Island, but not by Sonder.

Caulerpa laetevirens Mont.

C. sedoides (R. Br.) J. Ag. Upright fronds 3 to 4 inches long.

C. tristicha J. Ag. The ramenta regularly tristichous, short and mucronate.
Probably the form attributed to C. cupressoides by Sonder.

Udotea orientalis A. and E. 8. Gepp. Not noted by Sonder, but recorded from
Cape Flattery, Dunk Island and Port Denison (A. and E. S. Gepp, Codiaceae of
the Siboga Expedition).

U. argentea Zan. Not noted by Sonder, but collected by Banfield at Dunk
Island, and also sent to the British Museum from Queensland by Bailey.

One example of each of the Udoteas but in each case a fine specimen with
typical frond filaments.

Neomeris dumetosa Lamour. A single frond growing on a fragment of
coral, but identical in structure with specimens in the Sydney Herbarium,
distributed by Harvey from the Friendly Islands. I do not know of any previous
record of this plant from Australia.t

Avrainvillea erecta (Berkeley) A. and BE. 8S. Gepp. = A. papuana Murray and
Boodle. A single frond, dark green, stipitate, fan shaped with finely crenulate-
lacerate edges. Stipes 10 mm. long, 15 mm. wide near the base; lamina 30 mm.
long, 45 mm. wide. The filaments are cylindrical and yellow by transmitted light,
with a diameter of 30 to 40 w and somewhat swollen extremities.

Known to range from Madras and Ceylon to New Guinea, but not previously
recorded from Australia. (See Plate xli, fig. 3.)

1 Since writing the above, I have received further clumps of fronds from Mr. Rainford.

fod

BY A. H. S. LUCAS. 557

An allied species, A. clavatiramea, has been described by A. and E. S. Gepp
from plants gathered by J. Bracebridge Wilson in Corio Bay (Port Phillip) and
at Port Phillip Heads.

Otherwise, the genus is not represented in Australia.

Halimeda. Mr. Rainford has forwarded four species: H. tuna (Ellis and
Soland.) Lamour; H. opuntia (L.) Lamour; H. incrassata (EHllis) Lamour;
H. polydactylis J. Ag.

Codium. Sonder had no Codia from Port Denison but two forms from
Rockingham Bay, which he ascribes to C0. tomentosum, “the smaller scarcely two
inches, the larger in contrast over a foot long”.

Mr. Rainford also sends me two forms from Bowen; the smaller, a little over
two inches long and compressed, appears to agree in characters with C. lineare
J. Ag., previously recorded from Port Denison and from Moreton Bay; the larger,
about six inches long and terete, agrees with CO. tomentosum (Huds.) Stackh. The
utricles of both are obovate-pyriform with very thin apical membrane.

3. Two forms of Codium.

Mr. Rainford sent me a series of examples of a Codium, which has puzzled me.
The first two specimens were saccate and of a very dark green, and I thought
that I might be handling specimens of C. ovale Zan., described from New Guinea
as follows:

“Rronde pumila, substipitata, obovata, intus cava, intense colorata, exsiccatione
atrovirente’”. (De Toni, Sylloge Algarum, ii).

Plants sent later by Mr. Rainford were much more varied in form, some
resembling the irregular dense-textured and closely adhering plants of
C. adhaerens, which is common on our southern coasts, and others growing out
into ultimately hollow lobes.

Amongst the Codia of the Siboga Expedition collections from the East Indian
seas, A. and E. S. Gepp give C. adhaerens, C. difforme and C. ovale. De Toni in
the Sylloge includes C. difforme Kuetz. under C. adhaerens. Mr. and Mrs. Gepp
evidently regard C. difforme as a distinct form. We have no copy of Kuetzing’s
work in Australia, and I have had no opportunity of seeing his description. I had
drawn up a description of the Bowen plant, considering it to be distinct from
C. adhaerens, and I append the description. After reading the Siboga Monograph,
however, I suspect that the plant may very well be C. difforme, and that it bridges
the gap between C. adhaerens and C. ovale. The former grows between tides
and the latter is recorded at 16 m. from North Ubian, Sulu Archipelago, and at
27 m. from Banda. (Codiaceae of the Siboga Expedition.)

Frons polymorpha varie lobata vel tuberiformis hine et illinc byssis ad saxa
adfixa; cutis epidermea ex ultimis utriculis parallelis constans firma, fila interiora
laxa mucosa, utriculi ultimi cylindracei obtusi rarius subpyriformes, 75-108 wu lati,
plerumque 350-380 w longi; frons matura intus cava; substantia firma, fronde ad
atmosphaeram exposita vel in aqua dulci immersa haud faciliter deliquescente;
color obscure viridis, in sicco nigrescens. Sporangia non visa.

Habitat.—Bowen, Queensland (E. H. Rainford). (See Plate xli, figs. 1 and 2.)

I take the opportunity of describing here a form of Codium, which I gathered

in 1909 at Caloundra, South Queensland, but which was not forwarded to me
by Mr. Rainford.

558 NOTES ON AUSTRALIAN MARINE ALGAE, V,

CoDIUM GLOBOSUM, n. sp. (Plate xli, fig. 4.)

Frons globosa solida, bysso filorum sericeorum adfixa, 10-20 mm. diametro,
intus plexu filorum intricato, extus lobis periphericis inflato-cylindraceis ad
superficiem frondis directis, apicibus liberis eminentibus, constans. Fila interiora
cire. 30 uw lata, insuper viridia, flexuosa nunc dichotoma. Lobi terminales singulo
peripherico strato appositi, circ. 3 mm. longi, 275-350 uw lati, valde obtusi, contentum
viridem exhibentes, e filis singulis vel pluribus surgentes. Sporangia non visa.
Color laetevirens.

Habitat.—Caloundra, S. Queensland (Lucas).

Frond globose solid, attached by a byssus of silky threads, 10-20 mm. in
diameter, consisting of an inner intricate plexus of fibres, and externally of
peripheric inflated-cylindrical utricles directed to the surface, the apices freely
projecting. The interior fibres about 30 mw wide, flexuose and now and then
dichotomous, green in their upper range. Utricles adpressed in a single peripheric
stratum, about 3 mm. long and 275-350 wu broad, bluntly obtuse with a green
content, arising by done or more branches from the fibres. Sporangia not seen.
Colour light green.

Habitat—Caloundra, S. Queensland (A. H. S. Lucas).

A small form having the general shape and habit of C. bursa and
C. mammillosum, and apparently intermediate in characters between these two
species. The main distinction lies in the dimensions of the ultimate green lobules.
The diameter in the three species has the width as follows:

C. bursa from Victoria and Tasmania 120-200 wu.

C. globosum from S. Queensland 275-350 uw.

C. mammillosum from Warrnambool 600-770 wu.

The surface texture shows a corresponding difference in the three forms,
close in CG. bursa, more openly papillate in C. globosum and much more coarsely
mammillate in 0. mammillosum. The colour of the two latter is a brighter and
lighter green than that of C. bursa. (See Plate xli, fig. 4.)

4. Notes on Caulerpa.

In the Herbarium of the Botanic Gardens, Sydney, were examples of a
Caulerpa, which appeared to me to be C. ligulata Harv., recorded only from South
Africa. The specimens had been obtained from the herbarium of R. Helms, and no
locality was indicated. Later, however, I came across the plant growing abundantly
on mud-covered rocks at Sandringham, Botany Bay; and still later I found many
large plants washed up at Balmoral, Middle Harbour, Sydney. They agree well
with Harvey’s description, attaining a height of over a foot and forming stout
clumps with matted surculi spreading over an extensive base of several square
inches. The colour is a most vivid green. This is a very remarkable distribution
of a Caulerpa, a member of a genus in which reproduction by means of spores is
quite unknown, and throws an unexpected light on former land connections. (See
Plate xlii.)

Mr. R. Baxter sent a large collection of Algae from Lord Howe Island to the
Australian Museum, Sydney, which by the courtesy of the Director, Dr. Charles
Anderson, I was enabled to examine. Amongst them were fine specimens of
Caulerpa fastigiata Mont. This makes an addition to the Australian Caulerpa
flora, which now includes 41 species. We may anticipate that other Caulerpas

of the Pacific Islands will also be found on our tropical shores and reefs. ( See
Plate xliii, fig. 1.)

BY A. H. S. LUCAS. 559

In 1909 the collections of Algae made on an extensive voyage of the Govern-
ment trawler “Endeavour” were submitted to me. Mr. Charles Hedley accom-
panied the expedition and to his judgment and care the value of the algal
collection was due. When off Kangaroo Island, a Caulerpa, represented by four
or five specimens, was dredged in some eight fathoms of water. The plant was
evidently quite new to Australia, and I proposed to describe it under the name
of C. Hedleyi from the discoverer. To make sure that it was an altogether
unknown form, I sent a specimen to Mrs. E. S. Gepp of the British Museum,
who had kindly assisted me on other occasions. Mrs. Gepp forwarded it to
Madame Weber van Bosse in Belgium, who was working on Caulerpa at the time.
The latter then described the plant as new, and I am glad to say adopted the
name C. Hedleyi.

I append the description, which was published in the Annales de V’Institut
Océanographique (Prince de Monaco) ii, 1, pp. 1, 2, 1910, as this publication is not
generally available in Australia.

CAULERPA HrpLEYI A. Weber van Bosse. (Plate xliii, fig. 2.)

Frondibus constantibus surculo repenti, squamuloso, unde truncus ramosus
surgit. Squamuli identidem dichotomi, fere quadripartiti, apicibus bifurcis et
mucronatis surculum cingunt ordinibus compluribus, densatis.

Truncis ramis oppositis, patentibus; truncus et rami ramulosi sunt. Ramuli
identidem dichotomi sunt, adpressi, 500-600 u longi, apicibus bifurcis et mucronatis,
circum truncum et ramos verticillis irregularibus constantibus densatis positi.

Le (. Hedleyi ressemble par sa ramification, savoir—un axe central portant
des branches opposées, distiques, tres serrées—aux Caulerpes de la section des
Araucaroideae (including the four Australian species, C. flexilis, C. hypnoides,
C. Muelleri and C. abies), et par la forme des squamules et des ramules, a celles
des Bryoideae, et dans cette section, dont les ramules entourent l’axe principal et
les branches, surtout a la forme tormentella du C. Webbiana ou au C. Pickeringii,
algues endogenes du Pacifique. (See Plate xliii, fig. 2.)

Madame van Bosse also alludes to the depth of the water in which C. Hedleyi
was growing. Most of our Caulerpas have been gathered in rock pools and in
channels through which the tides scour, but they probably extend into deeper
water, and flourish there more luxuriantly. At a depth of 4-6 fathoms in Port
Esperance, Tasmania, I dredged abundance of fine plants of C. trifaria, much
larger than those in the rock pools of Port Phillip Heads, and also dredged
C. Brownii at similar depth in the D’Entrecasteaux Channel, while Mr. L. Rodway
dredged C. cactoides at 20 fm. near Actaeon Island.

Edible Green Algae. In a collection forwarded by my friend, Dr. Olive Wood,
from Nukualofa, Tonga, were three plants, which she informs me are eaten by
the natives, Caulerpa clavifera, Codium tomentosum and C. difforme. Of the first
Dr. Wood writes: “No. 1 is a great delicacy, and especially appeals to the palate
of women during pregnancy, who frequently leave their homes inland, and live
with relatives in a seaside village, so as to be near the source of supply’.

Miss Minnie Reed in a paper on “The Economic Seaweeds of Hawaii and
their Food Value”, published in the Annual Report of the Hawaii Agricultural
Experiment Station, 1906, states that there are over 70 distinct species of algae
used for food by the Hawaiians. Two of these are Codium adhaerens and
C. tomentosum.

560 NOTES ON AUSTRALIAN MARINE ALGAE, V,

5. Fucoideae. Distribution Notes.

In Notes on Australian Marine Algae, i (Proc. Linn. Soc. N.S.W., 1913, xxxviii,

p. 51) I recorded Hormosira articulata (Forsk.) Zan. from Port Stephens, and

ventured to predict its occurrence further north. Since that time, 1913, I have

received specimens from New Caledonia (C. Dézarnaulds), and now from Bowen,
Port Denison (HE. H. Rainford). (See Plate xliv.)

In the same paper I suggested that the Haliseris of N.H. Queensland was
probably not H. Muelleri but H. acrostichoides. Mr. Rainford has sent me typical
plants of H. acrostichoides from Bowen. Mr. H. A. Longman sent me specimens
from Noosa Hd., and I gathered it at Sandgate, Moreton Bay, so that this seems
to be the commonly occurring Haliseris of the east coast of Australia.

J. G. Agardh, Spec. Sargass. Australiae, gives the range of Sargassum
cristatum J. Ag. as southern coasts of Australia (F. von Mueller, Watts, Collie).
The Mesdames Harrison and Waterhouse gathered most characteristic fruiting
specimens on the beach at Geraldton, W.A.

Sargassum verruculosum (Mert.) J. Ag. appears to be very shy of fruiting.
Acres of the bottom of many Tasmanian harbours and estuaries are covered
with this plant but I never found a specimen in fruit.

It is very noteworthy that the Sargassa met with round the coasts of
Tasmania are almost or quite confined to the subgenera Phyllotricha’ and
Arthrophycus, while the species of Husargassum abound in the warmer seas of
Western and North-eastern Australia.

6. Two New Species of Chondria.

Chondria is well represented in Australian seas. De Toni lists 11 species.
Certain of these seem to be very close counterparts of European and West Indian
species. Thus Harvey distributed Australian forms under the names of
C. dasyphylla, C. sedifolia and C. tenuissima (?). By later writers these identifi-
cations have not been accepted.

In the present paper I venture to add two new species, one from Bowen
closely allied to C. tenwissima, and the other from N. S. Wales closely allied to
C. caerulescens. The former belongs to the section Huchondria, with the punctum
vegetationis freely produced at the tip of attenuate segments and the latter to the
section Coelochondria, with the punctum vegetationis immersed in a pit at the
apex of the blunt segments.

CHONDRIA RAINFORDI, n. sp. (Plate xlv, Plate xlvii, fig. 2.)

Frons robusta, plexu fibroso radicata caespitosa, pedalis et ultra, alterne
decomposito-pinnata; caulis teres vix compressus, a basi tenui gradatim superne
incrassatus, regione media pennam columbae crassitie aequans; rami alterni ad
20 uncias longi, 12-25 mm. distantils quoquoversum egredientes, ambitu lanceolati
vel fere lineares; rami ramulique (pinnae) utrinque conspicue attenuati; pinnae
superne bi- vel tripinnata copiosas pinnellas fere ad instar scoparum gerentes;
pinnellae lineares utriuque attenuatae, 4-7 mm. longae, longitudine 10-20 es
latitudinem excellente.

Substantia mollissima mucosa unde frons ad chartam tenacissime adhaeret.
Color purpurascens.

Cystocarpia subsessilia fructus Punicae-granati facies referentia, in parvis
inferioribus ramis modo inevolutis sedentia, sporas magnas pyriformes 150 u
longas foventia.

BY A. H. S. LUCAS. 561

Stichidia. ramulos ultimos pinnellae constituentia, sterilibus ramellis omnino
similia, vix torulosa nec tumida, sporangia supra mediam partem sparsa triangule
divisa gerentia.

Antheridia nondum visa.

Habitat—Bowen, Queensland.

Frond robust, caespitose, attached by a plexus of fibres, a foot or more high,
alternately decompound-pinnate; stem round, scarcely compressed, from a slender
base gradually thickened upwards, in the median region equalling a pigeon’s
plume in thickness; branches alternate, to 20 inches long, emerging in all
directions at intervals of 12-25 mm., lanceolate or almost linear in outline;
branches and branchlets (pinnae) conspicuously attenuated at both extremities;
pinnae bearing in their distal part copious bi-tripinnate almost brush-like pinnellae;
pinnellae linear attenuate at both extremities 4-7 mm. long, the length exceeding
ten times the breadth.

Substance very soft mucous so that the frond adheres most closely to the
paper.

Colour purpurascent.

Cystocarps subsessile like tiny pomegranates seated on small lower not greatly
developed branches, forming large pyriform spores 150 wu long.

Stichidia consisting in the ultimate divisions of a pinnella, altogether
resembling the sterile divisions, scarcely swollen or torulose, bearing triangularly
divided sporangia dispersed above the middle of the stichidium.

Antheridia not seen.

Habitat.—Bowen, Queensland.

I dedicate this elegant Chondria to Mr. EH. H. Rainford, of Bowen, who
gathered a fine sequence of specimens and mounted them while fresh. I am
greatly indebted to Mr. Rainford for excellent collections of algae from Port
Denison.

This is probably the plant recorded from Port Denison by Sonder under
the name C. tenuwissima J. Ag. He states that the plants were 1-13 feet high
(Die Algen des tropischen Australiens, p. 51).

By way of comparison I give photographs of C. fusifolia H. and H., which
I gathered in the Tamar and in the Derwent. The plants are crowded with
tetrasporangia, but I have not found any bearing cystocarps. (Plate xlvi, Plate
xlvii, fig. 1.)

CHONDRIA IRIDESCENS, n. sp. (Plate xlviii.)

Frons teres, plexu fibroso radicata, pyramidatim decomposito-pinnata, ad
25 cm. alta; rami copiosi, distantiis 5-10 mm. orti, quoquoversum vagi, ad 15 cm.
longi, pyramidatim evoluti, apices versus acuminati; ramuli crebrissimi, alterni,
quoquoversum crescentes, distantiis 3-4 mm. egredientes, plures aequaliter ad
12 mm. longi, ramellos pinnatim gerentes; ramelli cylindracei, ad 2 mm. longi,
fovea apicali obtusi. Rachides flexuosae. Substantia gelatinoso-carnea ergo
plantu chartae tenacius adhaeret. Color obscure purpureus, in vivo pulchre
coeruleo-iridescens. ,

Cystocarpia, Antheridia, nondum visa.

Stichidia ramellis conformia, subtorulosa, sporangia praecipue supra medium
foventia triangule divisa.

Sub lente cellulae superficiales ramellorum diametro aequales, ramulorum et
ramorum plures longiores apparent.

Habitat.—N. S. Wales (Port Hacking, Botany Bay, Port Stephens).

562 NOTES ON AUSTRALIAN MARINE ALGAE, V,

Frond round, attached by a fibrous plexus, pyramidate decompound pinnate,
to 25 em. high, branches copious, at distances of 5-10 mm., spreading in all
directions, pyramidate, acuminate towards the apices; secondary branches crowded,
alternate, at distances 3-4 mm., growing out in all directions, the greater part
of the middle ones equal, to 12 mm. long, bearing ramelli pinnately; ramelli
cylindraceous to 2 mm. long, obtuse with terminal hollow pit. Rachides flexuose.

Substance gelatinous fleshy so that the plant very closely adheres to paper.

Colour dark purple, when growing beautifully bluish iridescent.

Cystocarps and Antheridia not seen.

Stichidia quite like the sterile ramelli, subtorulose, bearing triangularly
divided sporangia chiefly above the middle.

Under the microscope the superficial cells of the ramelli are as broad as long,
those of the ramuli and rami many times as long as broad.

Habitat—N. S. Wales (Port Hacking, Botany Bay, Port Stephens).

This form is clearly very nearly related to C. coerulescens but of larger
dimensions. I examined specimens of C. curdieana Hary. in the Melbourne
Herbarium but the aspect of these was quite unlike that of the N.S.W. plants.

EXPLANATION OF PLATES XLI-XLVIII.
Plate xli.
1 and 2. Forms of Codium difforme? Bowen, Qld. (Rainford.)

3. Avrainvillea erecta = A. papuana. Bowen. (Rainford.)
4. Codium globosum, n. sp. Caloundra, Qld. (Lucas.)

Plate xlii.
Caulerpa ligulata. Balmoral, Middle Harbour, Port Jackson. (Lucas.)

Plate xliii.
1. Caulerpa fastigiata. Lord Howe Is. (R. Baxter.)
2. Caulerpa Hedleyi. Kangaroo Is. (C. Hedley.)

Plate xliv.
Hormosira articulata. Bowen, Qld. (Lucas.)

Plate xlv.
Chondria Rainfordi, n. sp. Bowen. (Rainford.)

Plate xlvi.
Chondria fusifolia. R. Tamar. Tas. (Lucas.)

Plate xlvii.
1. Chondria fusifolia. KR. Derwent, Tas. (Lucas.)
2. Chondria Rainfordi, n. sp. Base. Bowen (Rainford.)

Plate xlviii.
Chondria iridescens, n. sp. Port Hacking, N.S.W. (Kretschman.)

MOSQUITO CONTROL IN THE MUNICIPALITY OF LANE COVE,
“NEW SOUTH WALES.

By B. BERTRAM.
(Communicated by Dr. I. M. Mackerras.)

(Plates xlix and 1 and one Text-figure.)
[Read 30th November, 1927.]

Introduction.

Mosquito control may be instituted as a prophylactic measure against disease
or to abate a nuisance. While there does not appear to be any serious danger
from mosquito borne diseases in the area under consideration, mosquitoes were
sufficiently abundant to be a public nuisance. A survey was therefore carried out
in 1925 and a report, with recommendations, presented to the Lane Cove Council,
which authorized a small expenditure enabling control work to be carried out over
a limited area. The results obtained were so satisfactory that adequate provision
is now made for maintaining this work and extending it to other parts of the
Municipality.

Mosquito surveys have been carried out in various parts of New South Wales
by Ferguson (1922, 1927a) and Mackerras (1926), the last named dealing with
the Sydney district as a whole. Control work has also been undertaken more or
less thoroughly in several areas, of which Kyogle may be particularly mentioned.
This town had in the recent epidemic a dengue fever incidence of 10% as compared
with 90% for neighbouring towns in which no control had been attempted
(Ferguson, 19270). No description of the methods adopted or the results obtained
have, however, been published. Since the local problem has presented some
interesting features, it seems desirable that the results of the present investigations
should be made available.

I was indebted to the late Dr. Eustace W. Ferguson for determinations and
other assistance during the early part of the work.

Topography.

The Municipality of Lane Cove is a suburb of Sydney situated on the northern
slopes of Port Jackson. Though only four square miles in extent, it comprises
a variety of topographical features. It is bounded on the south and west by the
waters of Port Jackson and the Lane Cove River and includes high wind-swept
ridges of shale formation, and deeply dissected, shaded sandstone gullies, along
which flow permanent creeks, some direct into open bays in the Harbour and
some in salt water estuaries fringed with mangroves and headed by reedy brackish
swamps.

There is always some flow of water in these creeks even during the driest
weather, but in many places they consist of chains of rock pools or small ponds
dammed back by sand bars or debris. The foreshores consist partly of precipitous

s

564 MOSQUITO CONTROL,

sandstone cliffs fringed by saline rock pools and partly of mangrove covered mud
flats and in some places the low lying rock foreshores slope gently back towards
the ridges.

The more elevated parts of the area are more or less densely settled, and only
a small section is sewered, namely the St. Leonards area and portion of the
Greenwich area. Over the remainder of the area the greater part of the house
drainage eventually flows into the creeks, which are usually set aside as drainage
reserves in most subdivided land. This household effluent causes a considerable
pollution of the creeks, which extends for a variable distance down stream until
removed by aeration, filtration, and dilution by clear water tributaries and the
addition of subsoil water. The extent of this pollution, which is the most important
factor in the mosquito problem, is shown by cross lines in Text-fig. 1. The
extension of sewerage facilities is gradually reducing the amount of pollution,
while the institution of pipe drainage has eliminated the pollution in certain
areas. Unfortunately, however, the trouble caused by the presence of organic
pollution is not entirely eliminated by the institution of pipe drainage, though
it may transfer the point of outlet of the drainage to a more remote and less
settled area except in cases where the pipe line terminates in tidal waters.

LANE COVE

iy Tyo

“ZANE COVE

7. Gc '
AA7B0 URI: HAE er,
oA Bae Oro.

1
hota

Text-fig. 1. Map of the Municipality of Lane Cove. The area of denser
settlement is stippled, the points of pollution of the creeks are shown by circles,
and the extent of pollution by small cross lines.

BY B. BERTRAM. 565

Results of the Survey.
The following ten mosquitoes have been found in the district:

Culex fatigans Wiedemann.—This species was found to be a serious pest in
the densely settled areas, frequenting the dwellings in and around the business
and shopping centres to the detriment and inconvenience of the residents. The
factors governing the prevalence of this species are dealt with fully below.

Aedes (Ochlerotatus) vigilax Skuse.—Specimens were taken in the bush
gullies. It does not occur in the vicinity of dwellings in sufficient numbers to
warrant attention. It is not uncommon in the bush, but certainly does not appear
to be nearly so prevalent as it is in other parts of the Sydney district.

Aedes (Ochlerotatus) flavifrons Skuse.—Only two specimens of this species
have been taken, one biting about midday in June and one in the late afternoon
in October, 1927, in bush gullies.

Aedes (Finlaya) alboannulatus Macquart.—Larvae have been taken in soakage
pools and creek ponds, and on rare occasions in slightly stagnant and polluted
water. The adult appears to be more numerous in the bush than Aedes vigilaz,
and it attacks viciously in such situations, but does not seem to be a pest in houses.

Aedes (Finlaya) queenslandis Strickland.—Adults were taken occasionally
in shaded gullies.

Aedes (Finlaya) notoscriptus Skuse.—Adults were taken occasionally in
houses. Larvae were first taken in a small collection of rain water in an artificial
container. This water became very warm with the sunlight and contained a
quantity of decaying gum tree leaves. Similar collections of water have been
found to favour the breeding of this species.

Aedes (Pseudoskusea) concolor Taylor.—Larvae were found in saline littoral
pools at Greenwich in water with a high temperature due to direct sunlight,
whilst there was also a crust of salt forming around the edges of the pools. It
does not frequent houses.

Lutzia halifaxi Theobald.—Larvae have been taken in clear rock pools in
the creeks, usually in company with larvae of Aedes alboannulatus, on which it
is a predator.

Culex (Neoculex) fergusoni Taylor.—Larvae were collected in a clear rock
pool in the bush near Fig Tree. It was very plentiful on the occasion of the first
visit. A few days later no larvae in any stages of development were found, but
larvae of Zygoptera had become numerous.

Anopheles annulipes Walker.—Adults were shy biters and rarely met with.
In fact on only one occasion was a specimen observed on the wing. Adults were
found trapped on an oil film in a small soakage pool. Larvae have been taken in
clear water, in small numbers, and in fairly large numbers in early summer in
more or less muddy pools in company with C. fatigans.

From these findings it was apparent that Culex fatigans was the only species
requiring serious attention and it was decided from the point of view of economy
and efficiency to limit all attempts at control strictly to this species. It was
further considered desirable that adequate control should be obtained on Council
lands before attempting any house to house campaign.

The problem was to deal with C. fatigans breeding in natural collections of
water such as in creeks and drainage easements rather than in artificial collections
of water.

The dominant factor determining the extent of the breeding of C. fatigans in
all creeks and drainage easements was found to be the extent of the pollution.

566 MOSQUITO CONTROL,

In clear unpolluted water numerous predatory aquatic insects were found and.
C. fatigans was absent, except at the very beginning of the season before predatory
life became abundant.

In only one instance, where natural waters were polluted, was it found
that larvae of C. fatigans were not present at some time during the season. In
this particular case a large dam used for manufacturing purposes received heavy
organic pollution from the waste water returning from the machines, and the
fact that kerosene oil, which is used largely in the manufacturing process, gained
access to the dam undoubtedly accounted for the absence of larvae of C. fatigans.

In all other instances, however, the drainage-polluted bodies of water in the
creeks and drainage easements were found to be prolific sources of the breeding
of C. fatigans, and as these drainage easements and creeks came directly under
the Council’s control the work of control under such conditions was decidedly
a Council matter.

After the measures of control instituted by the Council became properly
established, the only instance where mosquitoes were reported to be on the
increase, was found on investigation to be due to a septic tank effluent gaining
access to the surface of the ground, due to faulty rubble absorption drains. With
the reconstruction of the rubble drains, and the effectual disposal therein of the
septic tank effluent, no further complaints were received.

That the presence of pollution due to a high content of organic nitrogenous and
other waste products was found to be favourable to the breeding of C. fatigans, was
indicated by the presence in large numbers of these larvae in the effluent of the
septic tank previously mentioned. In that case the effluent was seen to be flowing
over the ground surface for about 30 yards with a width of about 4 feet,
terminating in a grass covered area of several square yards. Here small pools
had been formed and each pool was found to contain hundreds of larvae of
C. fatigans.

Naturally such breeding areas would contain no aquatic predators, and there
is further evidence to indicate that both the type of nutriment and the absence
of predators determine the selection of breeding grounds by this species. This
was clearly demonstrated in one case, where three large ponds in a disused quarry
were inspected at various intervals during the season. No mosquito larvae were
found until towards the end of the season, when a rain storm carried a certain
amount of house drainage into one pond, and when visited about a week
later, this particular pond was found to contain enormous numbers of larvae of
C. fatigans, whilst the other unpolluted ponds remained free from mosquito larvae.

METHODS OF CONTROL.
Oiling.

Oil spraying was carried out only in correlation with the extent of pollution;
that is from the point of origin of pollution to the point where the presence of
gill breathing animals or insects indicated the elimination of organic putrescible
matter. In some cases this line of demarcation was fairly definite, as for instance
where at the end of a run of several hundred yards the water filtered through a
large natural sand filter in the form of a sand bar, or where a small drainage
easement entered a fast moving stream with a greater volume of water. Some-
times, however, it was not so definite, but the operator soon became skilled in the
work of oiling in such a manner as to produce the best results with the least
amount of damage to useful or harmless forms of life.

BY B. BERTRAM. 567

Plant.

The plant provided at the commencement of the season was rather obsolete,
and for want of staff the maintenance staff were given the work of oiling. Even
under such conditions favourable results were obtained, and at the commence-
ment of the following season two knapsack sprays and a horse and vehicle (the
latter in place of the wheelbarrow formerly used to transport equipment and oil)
were provided. The entire work was then given to one man engaged as a weekly
hand, his sole duties being that of mosquito eradication.

The results from then on were very satisfactory and mosquito control is now
looked upon by the Council as part of its ordinary duties, the Council no longer
allowing mosquitoes to breed on any country over which it has direct control.

This has placed the Council in the happy position of being able to guide and
direct the residents and ratepayers to take the necessary steps to rid their
premises of mosquitoes without themselves being the subject of adverse comment.

Channelling and Clearing.

The termination of the oil spraying at the end of last summer did not end
the Council’s activities in the work of mosquito control. As the result of a
recommendation the same weekly hand previously engaged on the spraying work
was employed on the work of channelling and clearing the drainage easements
and watercourses on which summer control measures had been adopted. It is a
well-known fact that C. fatigans seeks fairly still or sluggish water, and is also
very partial to shaded situations for breeding purposes, and, while the clearing
and channelling work will undoubtedly tend towards greater efficiency in mosquito
control during the coming season, it is also a sanitation measure which has long
been overlooked. Ponds of polluted water have been eliminated whenever possible
by channel cutting and in some localities rock channels have been made by blasting
with gelignite. Logs and debris have been removed and drainage polluted swamps
eliminated. From observations made during the following twelve months on
channels cut during the summer of 1926, there is every indication that a well cut
channel will require very little maintenance in this district besides greatly
reducing the expenditure on oil (Plate l, fig. 4).

Educational.

With a view of enlisting the co-operation of the residents and in the interests
of propaganda work on mosquito control, the Council accepted a recommendation
to provide the necessary equipment in the way of a projecting lantern and suitable
lantern slides to enable the writer to lecture to several local audiences. In this
Tespect every assistance and courtesy has been shown by the various head masters
of the schools and the leaders of other local organizations. The exhibition of
lantern slides at the local picture theatre free of charge has been due to the
interest and kindness shown by the proprietor to whom thanks are due.

Experiments with Automatic Oil Stations.

Drip oilers made from empty tins were found to be unsuitable, due to the
unwanted attention from the ubiquitous small boy. Bags of sawdust saturated with
oil, and submerged in the various drainage easements were not effective owing
to the excessive amount of solids and grease present in the drainage forming an
impervious scum around the bags.

T

568 MOSQUITO CONTKOL,

Laboratory Experiments.

Crude oil was found in laboratory experiments to be too heavy and flocculent,
allowing a clear area to form at the edge of the jar containing the larvae, and it
was also noticed that the larvae were capable of living beneath the oil film for
a period of three days. Close observation also showed that they were capable of
piercing the film, with their siphons, and on descending from the surface minute
pin holes were observed in the film.

An unrefined kerosene known as Solar oil produced efficient larvicidal results
in 13 minutes.

A mixture of equal parts of Solar oil and tar oil produced the desired
results in 7 minutes. The tar oil used without the Kerosene oil did not form as
complete a film as when the two oils were mixed.

As a result of these experiments the Council is now using a mixture of three
parts of unrefined kerosene and one part of tar oil in the field and the effects are
being carefully noted.

list of Former Prolific Breeding Grounds of Culex fatigans.

Phoenix Street at the outlet of drainage from Mills Estate was the first area
treated. Approximately 100 houses drain into the drainage easement at this point.
Here, prior to operations, the water immediately below the point of pollution was
dense with larvae, whilst a natural rock shelter near by was seen to be sheltering
myriads of the adult mosquitoes. (Plate xlix, fig. 2.)

From Little Street a drainage easement received drainage from the premises
on the eastern side of the shopping centre at the Lane Cove tram terminus, and
this easement for a distance of several hundred yards, together with two subsidiary
easements, formed another prolific source of mosquito breeding.

The drainage easements in the Helen Street-Norton Lane area were formerly
the origin of swarms of adult mosquitoes. (Plate 1, fig. 1.)

Tambourine Bay Creek required considerable attention for several hundred
yards down stream from the point of pollution. (Plate 1, fig. 3.)

Sofala Estate drainage which discharged into the Trouve Street drainage
easement, was effectually treated.

The Ronald Park Estate drainage easement, formerly ideal for mosquito
breeding, is now free from the pests.

Centennial Avenue and Barwon Creek have also been successfully treated,
whilst many other small unnamed drainage easements have received their quota
of expenditure.

Summary of the Work and its Results.

There is every indication that the work has been economically sound from a
Local Government expenditure point of view and will therefore be continued in
the future, whilst from a health and convenience standpoint there is every
indication that the residents have greatly benefited. In this respect one is
inclined to believe, judging by the benefit felt by residents at some considerable
distance from former breeding areas, that the range of flight of Oulex fatigans is
considerably greater than previously thought by workers in sanitary entomology.

BY B. BERTRAM. 569

EXPLANATION OF PLATES.
Plate xlix.

Pipe drain pouring polluted house drainage into a creek.

The pool to the left was heavily polluted and contained large numbers of larvae,
while myriads of adult C. fatigans were found sheltering under the overhanging
rock to the right of the picture.

3. A pool of still, clear water above the entry of the house drainage. No larvae of
C. fatigans were found here.

4. A small swamp in the course of a creek far enough below the point of pollution for the
water to have become again clarified. No larvae of C. fatigans were found here.

Plate 1.

1. A small heavily polluted pool in which large numbers of larvae of C. fatigans were
found.

2. The same pool as shown in Plate 1, fig. 1, after channelling and clearing had been
carried out.

3. A channel cut further down the creek where it runs in a sandstone gully.

An area which had been channelled and cleared twelve months before the photograph
was taken, during which time it had received no attention whatever.

References.

FERGUSON, E. W., 1922.—Notes on Insects. A.—Mosquitoes. Mosquito Survey of the
Murrumbidgee Irrigation Area. Rpt. Dir. Gen. Pub. Hlth. N.S.W. for Yr. 1920,
See. IV, 181-183.

———_—,, 1927a.— Mosquito Surveys of Some Inland Towns of New South Wales. Ibid.
for Yr. 1925, Sec. IV, 188-193.

,1927b.—The 1925-26 Outbreak of Dengue Fever in New South Wales. Ibid.
for Yr. 1926 (in press).

MackerraAs, I. M., 1926.—Mosquitoes of the Sydney District. Aust. Naturalist, vi, 3
33-42. '

A NEW DENDROBIUM FOR NEW SOUTH WALES AND QUEENSLAND.

By Rev. H. M. R. Rupp.
(Four Text-figures. )

[Read 30th November, 1927.]
DENDROBIUM TENUISSIMUM, 0D. Sp.

Planta semper pendula, cum ramis tenuissimis et lentis, 30-100 cm. longis.
Folia tereta, tenuissima, saepe brevia, obscure striata. Flores solae vel 2-3.
Sepala 14 cm. longa, lata, purpureo-fusca vel fusco-virida. Petala angusta, virida.
Labellum album, cum 5-6 maculis stipatis purpureis utrimque, et marginibus
crispis. Columna tenuis, apice purpureo.

An extremely slender, pendulous species occurring in dense brush-forests.
Stems almost like fine wire, very flexible, much branched. Leaves terete, in
young plants and branches somewhat robust, but becoming very slender in
maturity, occasionally as much as 15 cm. long, but more often very short, obscurely
fluted. Flowers solitary or in racemes of 2 or 3.. Sepals equal, about 1% cm.
long, rather broad, purplish-brown, or dark-green on the inner surface. Petals
shorter and very narrow, green. Labellum nearly as long as the sepals, recurved,
white with a yellowish-green median raised line, on either side of which are 4-6
closely-packed oval bright-purple blotches. Margins crisped-undulate, pure white.
Column slender, its apex and the stigmatic plate purplish. Ovary rather long,
pedicel long and exceedingly siender. Fl. September-October.

This plant has obvious affinities with D. Becklerii F.v.M., and D. Mortii F.v.M.,
but could never be mistaken for either. It is far more slender than even the less
robust forms of D. Becklerii, and never has the erect or semi-erect habit of that
species; the sepals are relatively much broader, the petals shorter : the labellum
is more obscurely lobed, and its margins are never bordered with purple, while the
brilliant purple blotches closely set on either side of the median plate are never
found in D. Becklerii. It differs from D. Mortii (which flowers in late summer
and autumn) in the slenderness and flexibility of the much longer branches; the
absence of angularity in the leaves; the relative dimensions of sepals and petals
(the latter in D. Mortii being very broad); and the brilliant purple markings of
the labellum. The ovary and pedicel in the new species are both relatively longer
than in either of the others. The perfume is very distinctive, and unlike that of
any other terete-leaved Dendrobium known to me. It is not particularly pleasant,
and has considerable resemblance to that of the common little liliaceous plant
Anguillaria dioica. .

Habitat—Upper Allyn River, Oct., 1925 (John Hopson and H.M.R.R.); Upper
Williams River, Oct., 1926 (C. Barrett); Upper Paterson River, 1927 (P. Laney) ;
Tambourine Mountain, Queensland, Sept., 1927 (Mrs. H. Curtis).

This orchid is found throughout the brushes among the foothills of the
Barrington Tops in New South Wales, the three rivers all having their origin in
that range. As it occurs also in South Queensland, it may be confidently looked
for in between. :

Text-fig.
Text-fig.
Text-fig.
Text-fig.

Be ow DH

ol
me)
=

bY H. M. R RUPP.

Dendrobium tenuissimum, n. sp., small flowering branch, about natural size.
Enlarged front view of flower of D. tenuissimum.

The same of D. Becklerii.

The same of D. Mortii.

Bet

ec) es eae
pM i Nh c
y bel vy

‘ i i
.~ 3 a A y ‘he len
: t i in?
e i ? , eye
1 i i % , uP
2 i ey i ye
i , aN NT yy,
il : ;
x : Pa i H ; )
ape ae j
i Lhe a j sey
f a! * ia i ri { a ib
! i oa P| ‘ ‘ 4}
i - i é ay, i
, i i ”
-

see |
oats

yp

IE ay aes Wak
es i 5 ; Sere,
(' :' ‘al i vat Pe i

th | it
i i ae
4 i
; Oy ee iy
i , ee
' ' vue
i UR
: i ‘ Lay ,
ae Mi
mY x
ig Fit
i is - Linh
‘ i
‘ i
vy | ’
i i
i) Pall i
. fe
‘
i a i ar: ; 1 '
\ ? 7
i il
mi i :
oy ‘ :
‘
rer y Uf
i
a J '
i < ; i
H i :
aa Ten

Proc. Linn. Soc. N.S.W., 1927. PLATE XXvii.

Jj Ps) o ee:

ra nidlip Heads. Western Crt
RAACEBRIDGE WILSON.

Collegted bys

A - eee 3 2 189: f-

Spongoclonium fasciculatum.

Lele mae

st

Proc. Linn. Soc. N.S.W., 1927. PLAVE XXvViii.

- Genus S P Pas ve lie ann

Species (eee. peer lata
Locality—Port Phillip Heads. S/” ‘ 7:
Collected by J. BRACEBRIDGE WILSON

Date 2 gp. ke 189 S

Spongoclonium brounianum from Victoria.

Aa

Varies.

PLATE XxXix.

Proc. Linn. Soc. N.S.W., 1927.

¥. wl2fay
VY foley

Rea 77. 4uy,

Spongoclonium brounianum from Western Australia.

Ge ssl
22 Ne

oat
ere
iy

Proc. Linn. Soc. N.S.W., 1927.

PLATE XXX.

3cm

Genus Callfo ae
Species wits v7 a); 4g.

Locality—Port Phillip Heads. SVahkisn 10
Collected by J. BRACEBRIDGE WILSON.
Datevo: /f 189 4.

Spongoclonrium wilsonianum.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXxXi.

la ©

Spongoclonium dasyurum.

Proc. Linn. Soc. N.S.W., 1927.

Spongoclonium paradoxum.

PLATE

PLATE XxXxiii.

Proc. Linn. Soc. N.S.W., 1927.

3em

Spongoclonium latissimum.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXXiv.

1

& 2 Mth PS hae enn gee tate Sra. +

Georg? Coe , Om 9 pS AT
Z Gy. OF Alor tg.

1. Spongoclonium angustatum.
2. Spongoclonium violaceum.

a.

Proc. Linn. Soc. N.S.W., 1927. ; E ; PLATE XXXV.

Sem

CLE Fac VEG

—~

4 ae ath, bay bree Cpr Lovie

O52 7 GO on

\

1. Callithamnion debile.
2. Callithamnion crispulwm.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXXVi.

Dampiera stricta R.Br.

Proc. Linn. Soc. N.S.W., 1927. PLATE XXXvii.

3-5. Style of Dampiera stricta in various stages.
6. Microspores germinating. j

ie
eter oily
es

ie

Proc. Linn. Soc. N.S.W., 1927. PLATE XxXxXvViii.

ae DAs ON CS:

1. Omphaloplacental region of a two weeks old embryo.
2. The downgrowth of mesoderm into the _ yolk-sac.

irs,

as

Proc. Linn. Soc. N.S.W., 1927. IZING 9-9'0-ab.<,

3. Yolk-sac of six weeks old embryo.
4. The mature allantoplacental region.
5. Folded allantoplacenta in one of the embryos examined.

PPATEH ox:

Proc. Linn. Soc. N.S.W., 1927.

«te

b

tion of inner and outer allantoic membranes in placental region.

Yolk-sac of eleven weeks old embryo.

6.

june

The

7.

Proc. Linn. Soc. N.S.W., 1927. PLATE Xi.

1 and 2. Forms of Codium difforme?

9
oO.

Avrainvillea erecta.
4. Codium globosum, n. sp.

Proc. Linn. Soc. N.S.W., 1927. PLATE xlii.

March 946 / Wd lo I

Caulerpa ligulata.

Proc. Linn. Soc. N.S.W., 1927. : PLATE xiii.

1. Caulerpa fastigiata. 2. Caulerpa Iedleyi.

Proc. Linn. Soc. N.S.W., 1927. PLATE xliv.

rapt tt

ae

Hormosira articulata.

py

ge oe
RS Re

Proc. Linn. Soc. N.S.W., 1927. PLATE xly.

BY tia

OM Mnf evel

Chondria Rainfordi, n. sp.

lak
\

FACE
ie

PLATE xlIvi.

Soc. N.S.W., 1927.

Proc. LINN.

Chondria fusifolia.

oe I Btn
: eas 2
:

‘a:

Proc. Linn. Soc. N.S.W., 1927. PLATE xlVvii.

anion 2
al

1. Chondria fusifolia. 2. Chondria Rainfordi, n. sp.

Proc. Linn. Soc. N.S.W., 1927. PLATE xviii.

Lal backng NI:

Chondria iridescens, n. sp.

EE TAY:
teh ke

i
Ar

Proc. Linn. Soc. N.S.W., 1927. . PLATE xlix.

Views taken in the Municipality of Lane Cove.

hee ote

Dy

ive We

athe
“ss genes
2

Proc. Linn. Soc. N.S.W., 1927. PrAnE lk

Views taken in the Municipality of Lane Cove.

bath
be

PRU

ot: ia

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fi ar i 5 .
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Notes on Australian Diptera. No. x. By Neat a Malloch.

by Dr. E. W. Ferguson.) (Twelve Text-figures.) woe Ne ae
The Influence of certain Colloids upon Fermentation. Parts JVVi... By
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Text-figures.) | :
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Weekes, B.Sc., Linnean Macleay Fellow of the Pee in Zoology.

“Connunien a ‘Sark

(Two

“By a Claire |

(Three Text-figures. ) aS as, ee 2
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Anophelini of the Mainland. By I. M. Mackerras, M-B., Ch.M. B.Sc, = :
Linnean Macleay Fellow of the Society in Zoology. eres: ene fae
figures. ) tie: ae 41 z
The Interpretation of fhe Radial Field of the Wing. in ‘the “Nematocerous - she rhe ;
Diptera, with Special Reference to the Tipulidae. By Charles 12s 3
Alexander, Ph.D. (Communicated by Dr. E. Ww. PEGA) Nincti ae Ge
two Text-figures.) 5o Se ae the See ee ne a eee ae AD 12,
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The Geology of the Country between Lamb’s Valley and the Paterson = ie
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New Gall-forming Thysanoptera of NGS tHALAS
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figures. ) : mane giana

An Ecological Study of fie Piers “of ANtotnt Wilson Bae iv.
Factors and Plant Response. By John McLuckie, M.A., D.Se., and

(Plate ix and fourteen Text-

Arthur H. K. Petrie, M.Sc. (Twenty Text-figures. ) Rey ere or
Two New Species of Setaria from Western Australia. By Dr. A. S.
Hitchcock. (Communicated by W. M. Carne.) Able Ne yom

By Dudley Moulton. —

Habitat

133-152. ir:
153-160

161-188; ees

185-186

_ (Issued 25th October, 1927.)

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By William Macleay, F.L.S. [1881].

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a

g

II

Notes. on Aastra Diptera No: xiii, P Jacl
by Dr. I. M. Mackerras.) (Fifteen, "Texas ur s.)
Notes on Australian. and Exotic ao :

Genus ‘Spongoctonium.
_ XXViL-xxv. ve) tes a :

(R. Br.) By P. praucle ware ]
and ieee ciets Text Reures) ©

: (Hinutia) quoyi. ‘By H. Claire Weekes, B. Se.,
of the Society in Zoology. oo xxxvilial 2

oe figures. ) eee et
-_Notes on Australian Marine “Alsae,_
{Plates xli-xlviii.) iss

Mosquito Control in the ‘Municipality of ‘Lané

. By B. Bertram. “(Communicated by Dr. 5 Bye
xlix-l, and one He Rene) ae os SARS

gies

_H. M. R. Rupp. (Four Text. figures. .

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mS

(Issued 15th February, 1928.)

THE

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|| LINNEAN SOCIETY
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“Abstract of Proceedings ..

_ PROCEED

Donations and Exchanges
List of Members .. .
Index epee aay Silecataadicg ate

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